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HARVARD UNIVERSITY 

Library of the 

Museum of 

Comparative Zoology 



The CANADIAN 
FIELD-NATURALIST 

Published by THE OTTAWA FIELD-NATURALISTS' CLUB, Ottawa, Canada 




Volume 104, Number 1 



January-March 1990 



The Ottawa Field-Naturalists' Club 

FOUNDED IN 1879 

Patron 

His Excellency The Right Honourable Ramon John Hnatyshyn, P.C, C.C, C.M.M., Q.C., 

Governor General of Canada 

The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada's natural 
heritage; to encourage investigation and publish the results of research in all fields of natural history and to diffuse 
information on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, 
maintaining or restoring environments of high quality for living things. 



Edward L. Bousfield 
Irwin M. Brodo 
William J. Cody 
William G. Dore 
R. Yorke Edwards 
Clarence Frankton 



Honorary Members 



Claude E. Garton 
W. Earl Godfrey 
C. Stuart Houston 
Louise de K. Lawrence 
Thomas H. Manning 
Don E. McAllister 



Stewart D. MacDonald Hugh M. Raup 



George H. McGee 
Verna Ross McGiffin 
Hue N. MacKenzie 
Eugene G. Munroe 
Robert W. Nero 



Loris S. Russell 
Douglas B. O. Savile 
Pauline Snure 
Mary E. Stuart 
Sheila Thomson 



1990 Council 



President: 
Vice-Presidents: 

Recording Secretary: 
Corresponding Secretary: 
Treasurer: 



Jeff Harrison 

Roy John 
Don Cuddy 

Elizabeth Fox 

Eileen Evens 
Mike Scromeda 



Ronald E. Bedford 
Barry Bendell 
Steve Blight 
William J. Cody 
Francis R. Cook 
Don Davidson 
Enid Frankton 
Deirdre Furlong 



Colin Gaskell 
Bill Gummer 
Paul Hamilton 
Elizabeth Morton 
Michael Murphy 
Frank Pope 
Kenneth Strang 
Doreen Watler 



LOP IJO 
La Roi, Department 



of Botany, 



Those wishing to communicate with the Club should address correspondence to: The Ottawa Field-Naturalists' Club, 
Box 3264, Postal Station C, Ottawa, Canada Kl Y 4J5. For information on Club activities telephone (613) 722-3050. 

The Canadian Field-Naturalist 

The Canadian Field- Naturalist is published quarterly by The Ottawa Field-Naturalists' Club. Opinions and ideas 
expressed in this journal do not necessarily reflect those of The Ottawa Field-Naturalists' Club or any other agency. 

Editor: Francis R. Cook, R. R. 3, North Augusta, Ontario KOG 1 RO; (6 1 3) 996- 1 755; Assistant to Editor: 

P. J. Narraway; Copy Editor: Louis L'Arrivee 
Business Manager: William J. Cody, Box 3264, Postal Station C, Ottawa, Ontario K1Y4J5 

(613)996-1665 
Book Review Editor: Dr. J. Wilson Eedy, R. R. 1, Moffat, Ontario 
Coordinator, The Biological Flora of Canada: Dr. George H. 

University of Alberta, Edmonton, Alberta T6G 2E9 
Associate Editors: Anthony J. Erskine 

C. D. Bird W. Earl Godfrey 

Brian W. Coad Charles Jonkel 

Diana Laubitz 
Chairman, Publications Committee: Ronald E. Bedford 

All manuscripts intended for publication should be addressed to the Editor at home address. 

Subscriptions and Membership 

Subscription rates for individuals are $20 per calendar year. Libraries and other institutions may subscribe at the rate 
of $35 per year (volume). The Ottawa Field-Naturalists' Club annual membership fee of $20 includes a subscription to 
The Canadian Field-Naturalist. All foreign subscribers (including USA) must add an additional $3.00 to cover 
postage. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should 
be* mailed to: The Ottawa Field-Naturalists' Club, Box 3264, Postal Station C, Ottawa, Canada KIY 4J5. 
Second Class Mail Registration No. 0527 Return Postage Guaranteed. 

Back Numbers and Index 

Most back numbers ol this journal and its predecessors. Transactions of The Ottawa Field-Naturalists' Cluh, 1879- 
1 886, and The Ottawa Naturalist. 1 887- 1919, and Transactions of The Ottawa Field- Naturalists ' Club and The Ottawa 
Naturalist ~ Index compiled by .lohn M. (iillctt, may be purchased from the Business Manager. 

C over: The Fourhorn Sculpin, M yoxocephalus quadricornis. Lady Franklin Point, Northwest Territories, July 1979. 
Photograph by R. K. Kashino of Vancouver, courtesy of D. E. McAllister, Canadian Museum of Nature, 
Ottawa. Sec status report by .L Houston, pages 7 13. 



W. Bruce McGillivray 
William O. Pruitt, Jr. 
Stephen M. Smith 
Constantinus G. Van Zyll de Jong 



MCZ 
LIBRARY 

OCT 3 1990 

HARVARD 
UNIVERSITY 



THE CANADIAN 
FIELD-NATURALIST 



Volume 104 
1990 



THE OTTAWA FIELD-NATURALISTS' CLUB 

Ottawa Canada 



The Canadian Field-Naturalist 

Volume 104, Number 1 January-March 1990 



Rare and Endangered Fishes and Marine Mammals of Canada: 
COSEWIC Fish and Marine Mammal Subcommittee 
Status Reports: VI 

R. R. Campbell 

Department of Fisheries and Oceans, 200 Kent Street, Ottawa, Ontario Kl A 0E6 

Campbell, R. R. Editor. 1990. Rare and endangered fishes and marine mammals of Canada: COSEWIC Fish and 
Marine Mammal Subcommittee Status Reports: VL Canadian Field-Naturalist 104(1): 1-6. 

Twenty-one status reports representing those species of fish and marine mammals which were assigned status at the 
1989 COSEWIC General Meeting have been prepared for publication. Committee and Subcommittee (Fish and 
Marine Mammals) activities are briefly discussed, including changes to category definitions introduced in 1988. Also 
of interest, is the creation of the RENEW Committee which will attempt to carry the process to the next logical step — 
recovery plans and implementation, for endangered species. Current lists of status assignment for fish and marine 
mammals and for species yet to be considered, or which are under consideration, are presented in tabular form. 

Vingt-et-un rapports de statut representant tout les donnees de statut aux poissons et aux mammiferes marins donnes 
un statut a la reunion de CSEMDC ont ete prepares pour publication. Les activites du Comite et du sous-comite 
(poissons et mammiferes marins) sont brievement discutees incluyant des changements aux definitions de categoric, 
introduitent en 1988. On presente aussi des renseignements concernant le Comite "RENEW" qui essayera de porter le 
processus a la prochaine etape logique — planification et implementation des planifications pour les especes en danger 
de disparition. Listes currents des especes aux poissons et aux mammiferes marins deja nominees et des especes 
lesquelles on a encore besoin de considerer ont montre en forme tabulaire. 

Key Words: Rare and endangered species, fish, marine mammals, COSEWIC, status, Canada. 

As indicated in previous submissions (Campbell able to use this journal as one step in achieving the 

1984, 1985, 1987, 1988 and 1989), the intent of the goal. A series of reports were published in 1984, 

Subcommittee on Fish and Marine Mammals is to 1985, 1987, 1988, and 1989 [see Canadian 

publish the status reports (on those species of fish Field-Naturalist 98(1): 63-133; 99(3): 404-450; 

and marine mammals) which the Committee on 102(2): 165-309; 103(2): 147-239] and the 

the Status of Endangered Wildlife in Canada encouraging response to these publications has 

(COSEWIC) have reviewed, approved and used as enabled us to continue. 

a basis for the assignment of status to species in Contributions to the Committee of $10 000 

jeopardy in Canada. The group of 21 reports made by the Department of Fisheries and Oceans 

presented herein represent the fish and marine and Environment Canada in 1988 were once again 

mammal component of those species assigned matched by World Wildlife Fund Canada, 

status in 1989. It is hoped that we will have the permitting the contracting of several new reports in 

continuing support of the Department of Fisheries 1988. The Fish and Marine Mammal Subcommit- 

and Oceans to offer, in succeeding volumes, those tee was able to take advantage of this and 25 

reports reviewed in future years (Table 1 presents reports were initiated in 1988-1989. Although there 

those species assigned status to April 1989). are still a number of reports in preparation or 

under review (Table 2), the number of species still 

Progress awaiting consideration has been reduced to 25 

COSEWIC has undertaken to make available to (Table 3). It is the Subcommittee's goal to clear this 

all Canadians, supporting information on each list by 1992. 

species classified (see Cook and Muir 1984). The There are currently 28 status reports on fish 

Fish and Marine Mammal Subcommittee has been species, and 26 on marine mammal species under 



The Canadian Field-Naturalist 



Vol. 104 



Table 1. Fish and marine mammal species with assigned COSEWIC status to April 1989. 



Species 



Scientific Name 



Status 



Date Assigned 



Fish 

Lake Sturgeon 

Bloater 

Blueback Herring 

Hornyhead Chub 

River Chub 

Redfin Shiner 

Golden Redhorse 

Least Darter 

River Darter 

Green Sunfish 

Longear Sunfish 

Spoonhead Sculpin 

Brook Silverside 

Lake Lamprey* 

Green Sturgeon 

Shortnose Sturgeon 

Spotted Gar 

Kiyi 

Squanga Whitefish* 

Pacific Sardine 

Silver Chub 

Umatilla Dace 

Bigmouth Shiner 

Pugnose Shiner 

Silver Shiner 

Pugnose Minnow 

Redside Dace 

Speckled Dace 

Central Stoneroller 

Banded Killifish (Newfoundland) 

Blackstripe Topminnow 

Bigmouth Buffalo 

Black Buffalo 

Spotted Sucker 

River Redhorse 

Brindled Madtom 

Orangespotted Sunfish 

Redbreast Sunfish 

Fourhorn Sculpin (Arctic Islands) 

Giant Stickleback* 

Unarmoured Stickleback* 

Blackline Prickleback 

Bering Wolffish 

Lake Simcoe Whitefish* 

Blackfin Cisco 

Shortnose Cisco 

Shortjaw Cisco 

Dcepwater Sculpin 

(Great Lakes Watershed) 
Black Redhorse 
Copper Redhorse* 
Margined Madtom 
Enos Lake Stickleback* 
Shorthcad Sculpin 
Aurora Trout 
Acadian Whitefish* 
Salish Sucker 
Gravel Chub 
Paddlcfish 
Dcepwater Cisco 



A cipenser fulvescens 


NIAC 


April 


986 


Coregonus hoyi 


NIAC 


April 


988 


Alosa aesiivalis 


NIAC 


April 


980 


Nocomis biguttatus 


NIAC 


April 


988 


Nocomis micropogon 


NIAC 


April 


988 


Notropis umbratilis 


NIAC 


April 


988 


Moxostoma erythrurum 


RANSDR^ 


April 


989 


Etheostoma microperca 


RANSDR 


April 


989 


Percina shumardi 


RANSDR 


April 


989 


Lepomis cyanellus 


NIAC 


April 


987 


Lepomis megalotis 


NIAC 


April 


987 


Cottus ricei 


RANSDR 


April 


989 


Labidesthes sicculus 


RANSDR 


April 


989 


Lampetra macrostoma 


Rare 


April 


986 


Acipenser medirostris 


Rare 


April 


987 


Acipenser brevirostrum 


Rare 


April 


980 


Lepisosteus oculatus 


Rare 


April 


983 


Coregonus kiyi 


Rare 


April 


987 


Coregonus sp. 


Rare 


April 


988 


Sardinops sagax 


Rare 


April 


987 


Hybopsis storeriana 


Rare 


April 


985 


Rhinichthys umatilla 


Rare 


April 


988 


Notropis dorsalis 


Rare 


April 


985 


Notropis anogenus 


Rare 


April 


985 


Notropis photogenis 


Rare 


April 


983** 


Notropis emiliae 


Rare 


April 


985 


Clinostomus elongatus 


Rare 


April 


987 


Rhinichthys osculus 


Rare 


April 


980+ 


Campostoma anomalum 


Rare 


April 


985 


Fundulus diaphanus 


Vulnerable'' 


April 


989 


Fundulus notatus 


Rare 


April 


985 


Ictiobus cyprinellus 


Vulnerable 


April 


989 


Ictiobus niger 


Vulnerable 


April 


989 


Minytrema melanops 


Rare 


April 


983 


Moxostoma carinatum 


Rare 


April 


983** 


Notorus miurus 


Rare 


April 


985 


Lepomis humilis 


Vulnerable 


April 


989 


Lepomis auritus 


Vulnerable 


April 


989 


Myoxocephalus quadricornis 


Vulnerable 


April 


989 


Gasterosteus sp. 


Rare 


April 


980 


Gasterosteus sp. 


Rare 


April 


983 


Acantholumpenus mackayi 


Vulnerable 


April 


989 


Anarichus orien talis 


Vulnerable 


April 


989 


Coregonus clupeaformis spp. 


Threatened 


April 


987 


Coregonus nigripinnis 


Threatened 


April 


988 


Coregonus reighardi 


Threatened 


April 


987 


Coregonus zenithicus 


Threatened 


April 


987 


Myoxocephalus thompsoni 


Threatened 


April 


987 


Moxostoma dusquesnei 


Threatened 


April 


988 


Moxostoma hubhsi 


Threatened 


April 


987 


Noturus insignis 


Threatened 


April 


989 


Gasterosteus sp. 


Threatened 


April 


988 


Cottus confusus 


Threatened 


Noven 


iber 1983 


Salvetinus fontinalis timagamiensis 


Endangered 


April 


987* 


Coregonus canadensis 


Endangered 


April 


983 


Catostmus sp. 


Endangered 


April 


986 


Hybopsis x-punctata 


Extirpated 


April 


987*** 


Polyodon spalhula 


Extirpate J 


April 


987 


Coregonus johannae 


Extinct 


April 


988 



1990 



CAMPBELL: FiSH AND MARINE MAMMAL STATUS REPORTS 



Table I. Fish and marine mammal species with assigned COSEWIC status to April 1989 (concluded). 



Species 


Scientific Name 


Status 


Date Assigned 


Fish (continued) 










Lonjaw Cisco 


Coregonus alepnae 


Extinct 


April 


1988 


Banff Longnose Dace* 


Rhinichthys cataractae smithi 


Extinct 


April 


1987 


Blue Walleye 


Stirzostedion vitreum glaucum 


Extinct 


April 


1985 


Marine Molluscs 










Northern Abalone 


Haliotis kamtschatkana 


N/A 


April 


1988 


Marine Mammals 










California Sea Lion 


Zalophus californianus 


NIAC 


April 


1987 


Stellar Sea Lion 


Eumetopias jubatus 


NIAC 


April 


1987 


Atlantic Walrus 


Odobenus rosmarus rosmarus 








Eastern Arctic 




NIAC 


April 


1987 


Northwest Atlantic 




Extirpated 


April 


1987 


Grey Whale 


Eschrichtius robustus 








Northeast Pacific 




NIAC 


April 


1987 


Northwest Atlantic 




Extirpated 


April 


1987 


Hooded Seal 


Cystophora cristata 


NIAC 


April 


1986 


Northern Elephant Seal 


Mirounga angustirostris 


NIAC 


April 


1986 


Ringed Seal 


Phoca hispida 


RANSDR 


April 


1989 


Dall's Porpoise 


Phocoenoides dalli 


RANSDR 


April 


1989 


Narwhal 


Monodon monoceros 


NIAC 


April 


1986** 


Blainville's Beaked Whale 


Mesoplodon densirostris 


RANSDR 


April 


1989 


Hubb's Beaked Whale 


Mesoplodon carlhubbsi 


RANSDR 


April 


1989 


Stejneger's Beaked Whale 


Mesoplodon stejnegeri 


RANSDR 


April 


1989 


True's Beaked Whale 


Mesoplodon mirus 


RANSDR 


April 


1989 


Beluga 


Delphinapterus leucas 








Beaufort Sea 




NIAC 


April 


1986 


St. Lawrence River 




Endangered 


April 


1983 


Eastern Hudson Bay 




Threatened' 


April 


1988 


Unagava Bay 




Endangered' 


April 


1988 


Sowerby's Beaked Whale 


Mesoplodon bidens 


Vulnerable 


April 


1989 


Blue Whale 


Balaenoptera musculus 


Rare 


April 


1983 


Fin Whale 


Balaenoptera physalus 


Rare 


April 


1987* 


Sea Otter 


Enhydra lutris 


Endangered 


May 


1978° 


Humpback Whale 


Megaptera novaeangliae 








Northeast Pacific 




Threatened 


April 


1982- 


Northwest Atlantic 




Rare 


April 


1985 


Bowhead Whale 


Balaena mysticetus 


Endangered 


April 


1980° 


Right Whale 


Eubalaena glacialis 


Endangered 


April 


1980/85^ 


Sea Mink 


Mustela macrodon 


Extinct 


April 


1985 



NIAC — Not in Any COSEWIC Category (i.e., not in jeopardy) 

N/A — Status not assigned. COSEWIC has no mandate for invertebrates. Report accepted and recommended NIAC 

Status agreed to, but not assigned. 
aRANSDR — Use of NIAC dropped in 1988 — RANSDR is not a category = report accepted no status designation 

required. 
*Endemic to Canada 
'"Vulnerable — 'Rare' category changed to 'Vulnerable' in 1988. Previous rare designations to be re-examined prior to 

automatic change to vulnerable. 
tUpdated April 1984 — no status change. 

"Updated April 1985 — North Atlantic stock downlisted to "Rare". 

'To be reviewed in 1990. 

"Updated April 1985 — no status change. 

"Updated April 1986 — no status change. 

**Updated April 1987 — no status change. 

***Updated April 1987 — previous status of 'Endangered' assigned April 1985. 



review or in preparation (Table 2); several will be 
assigned status in 1990. In addition to soliciting 
further status reports on species of concern, the 



Subcommittee continues to obtain updates on the 
status of selected species as new information 
becomes available. 



The Canadian Field-Naturalist 



Vol. 104 



Table 2. Fish and marine mammal species for which status reports are in preparation, or under review August 1989. 



Species 



Scientific Name 



Proposed Status 



Fish 

Chestnut Lamprey 
Darktail Lamprey 
Northern Brook Lamprey 
Atlantic Sturgeon 
Lake Sturgeon{ 
White Sturgeon 
Atlantic Salmon 
Bull Trout 
Bering Cisco 
Spring Cisco* 
Opeongo Whitefish* 
Pygmy Whitefish 
Pygmy Smelt 
Grass Pickerel 
Cutlips Minnow 
Eastern Silvery Minnow 
Ghost Shiner 
Lake Chubsucker 
Jasper Longnose Sucker* 
Mountain Sucker 
Leopard Dace 
Greenside Darter 
Warmouth 
Striped Bass 
Eastern Sand Darter 
Y-Prickleback 
Pixy Poacher* 
Bluefin Tuna 

Marine Mammals 

White-beaked Dolphin 
Harbour Porpoise 
Baird's Beaked Whale 
Cuvier's Beaked Whale 
Beluga Whale 

Cumberland Sound 

Western Hudson Bay 

Ungava Bay° 

Eastern Hudson Bay/ James Bay° 
Northern Bottlenose Whale 
Bowhead Whale° 
Killer Whale 
Long-finned Pilot Whale 
Right Whale° 
False Killer Whale 
Risso's Dolphin 
Sperm Whale 

Pacific White-sided Dolphin 
Northern Right Whale Dolphin 
Striped Dolphin 
Bottlenose Dolphin 
Short-finned Pilot Whale 
Pygmy Sperm Whale 
Dwarf Sperm Whale 
Atlantic White-sided Dolphin 
Common Dolphin 



Ichthyomyzon castaneus 


Vulerable 


Lethenteron alaskense 


Vulnerable 


Ichthyomyzon fossor 


Vulnerable 


Acipenser oxyrhynchus 


7 


Acipenser fulvescens 


7 


Acipenser transmontanus 


7 


Salmo salar 


7 


Salvelinus confluentus 


Vulnerable 


Coregonus laurettae 


7 


Coregonus sp. 


7 


Coregonus sp. 


Threatened 


Prosopium couheri 


7 


Osmerus spectrum 


Vulnerable 


Esox americanus vermiculatus 


Vulnerable 


Exoglossum maxillingua 


Vulnerable 


Hybognathus nuchalis regius 


Vulnerable 


Notropis buchanani 


Vulnerable 


Erimyzon sucetta 


Vulnerable 


Castostomus castostomus lacustris 


Vulnerable 


Castostomus platyrhynchus 


Vulnerable 


Rhinichthys falcatus 


7 


Etheostoma blennioides 


Vulnerable 


Lepomis gulosus 


Vulnerable 


Morone saxatilis 


Endangered 


Ammocrypta pellucida 


Vulnerable 


Allolumpenus hypochromus 


Vulnerable 


Occella impi 


Vulnerable 


Thunnus thynnus 


7 


Lagenorhynchus albirostris 


7 


Phocoena phocoena 


7 


Berardius bairdii 


7 


Ziphius cavirostris 


Vulnerable 


Delphinapterus leucas 






Threatened 

9 




Endangered 




Threatened 


Hyperoodon ampullata 


7 


Balaena mysticetus 


Endangered 


Orcinus orca 


7 


Globicephela malaena 


7 


Eubalaena glacialis 


Endangered 


Pseudorca crassidens 


7 


Grampus griseus 


7 


Physeter catadon 


7 


Lagenorhynchus obliquidens 


7 


Lissodelphis horealis 


7 


Stenella coeruleoalba 


7 


Tursiops truncatus 


7 


Globicephala macrorhynchus 


Vulnerable 


Kogia hreviceps 


Vulnerable 


Kogia simus 


Vulnerable 


Lagenorhynchus aculus 
Delnhinus delnhis 


7 

7 



*Endemic to Canada. 
° Updated Status Report. 



1990 



CAMPBELL: FiSH AND MARINE MAMMAL STATUS REPORTS 



Table 3. Fish and Marine Mammal Species of Interest to COSEWIC — August 1989 (Not listed by priority) 



Species 



Scientific Name 



Possible Status 



Fish 

Red (Arctic) Chari 



Lake Herring 



Lake Whitefish 



Mira Whitefish 
Round Whitefish 



Pygmy Longfin Smelt* 



Chain Pickerel 

Redfin Pickerel 

Chiselmouth 

Bluntnose Minnow 

Western Silvery Minnow 

Blackchin Shiner 

Rosyface Shiner 

Striped Shiner ' 

Weed Shiner 

Nooky Dace 

Liard Hotspring Lake Chub* 

Flathead Catfish 

Northern Madtom 

Texada Stickleback* 

Tessellated Darter 

Channel Darter 

Cultus Pygmy Coastrange Sculpin* 

Mottled Sculpin 

Spinynose Sculpin 



Salvelinus alpinus spp. 



Coregonus artedi 



Coregonus clupeaformis 



Coregonus sp 
Prosopium cylindraceum 



Spirinichus thaleichthys 



Esox niger 

Esox americanus americanus 
Acrocheilus alutaceus 
Pimphales notatus 
Hybognathus argyritis 
Notropis heterodon 
Notropis rubellus 
Notropis chrysocephalus 
Notropis texanus 
Rhinichthys cataractae spp. 
Couesius plumbeus spp. 

Pylodictis olivaris 
Noturus stigmosus 
Gasterosteus sp. 
Ethestoma olmstedi 
Percina copelandi 
Cottus aleuticus 
Cottus bairdi 
Asemichthys taylori 



? (Landlocked populations — 

Quebec, New Brunswick, 

Newfoundland/ Labrador) 
Endangered in Lakes Erie and 

Ontario but widespread 

elsewhere. 
Threatened in Lakes Erie and 

Ontario but widespread 

elsewhere. 
Vulnerable 
Vulnerable (Lakes Huron and 

Ontario but widespread 

elsewhere). 
Vulnerable (landlocked 

population in Harrington Lake, 

British Columbia) 
Vulnerable (Quebec, New Brunswick, 

Nova Scotia) 
Vulnerable (Quebec) 
Vulnerable (British Columbia) 
Vulnerable (Manitoba) 
? (Alberta) 

Vulnerable (Manitoba) 
Vulnerable (Manitoba) 
Vulnerable (Ontario) 
Vulnerable (Manitoba) 
Vulnerable (British Columbia) 
Vulnerable (British Columbia: Liard 

Hotspring) 
? Ontario 

Vulnerable (Ontario) 
Vulnerable 
Vulnerable (Ontario) 
Vulnerable (Ontario) 
Threatened (British Columbia) 
Vulnerable (British Columbia, Alberta) 
Vulnerable (British Columbia) 



*Endemic to Canada. 
'Not of immediate concern. 



I 



The 1989 General Meeting was also noteworthy 
in that changes in category definitions, introduced in 
1988 {see Campbell 1988) were used for the first time 
in assignment of status. The use of "vulnerable" in 
status assignment commenced in 1989. Species 
assigned a "rare" status prior to, and including, 
August 1988 will remain as designated for now, but 
Subcommittee chairmen have been requested to 
provide recommendations for adoption of a change 
from "rare" to "vulnerable" on the basis of an 
updated report where necessary, and the status 
reaffirmed or redesignated as appropriate at the 
1990 General Meeting. Species formerly declared to 
be "not in any category" and several listed as 
"threatened" may now require re -evaluation as well. 

Due to confusion arising from the use of the 



designation of "not in any COSEWIC category" 
(NIAC), the use of this terminology has been 
discontinued. Commencing in 1989, species for 
which no status is assigned are simply annotated 
"report accepted, no status designation required". 
This will be accompanied by a brief explanatory 
note to indicate reasons, i.e., populations are stable 
and of no concern at this time, therefore, no status 
designation is required. 

New Initiatives 

A new national strategy for the prevention of 
extinction of wildlife species in Canada was 
launched in 1988. Although not a part of 
COSEWIC, this initiative arose through the 
impetus of COSEWIC and is intended to carry the 



The Canadian Field-Naturalist 



Vol. 104 



process to the next logical step. This strategy, 
acronym "RENEW" (for Recovery of Nationally 
Endangered Wildlife) will use COSEWIC status 
assignments and reports as the basis for selecting 
species for attention and recovery plans. 

The concept is for all agencies, organizations, and 
individuals, interested and able to assist in the effort 
to preserve species, to agree to work together and 
follow the same process to achieve the same goal. 
The goal is to prevent species known to be at risk of 
extinction from becoming so, and to prevent other 
species from becoming at risk. The process will 
involve recovery programs for each species, based 
on comprehensive technical plans prepared by the 
best experts available. These will be approved by 
governments, and implemented in cooperation with 
other countries, individual interests, and other 
interested parties. 

It is beyond the scope of this editorial to give a 
detailed explanation of the strategy. Interested 
parties may receive further detailed information by 
contacting the Office of Primary Interest, RENEW, 
Canadian Wildlife Service, Environment Canada, 
Ottawa, Ontario KIA 0H3. 

Acknowledgments 

First and foremost, our thanks to the various 
authors who have so generously contributed their 
time and talent in support of COSEWIC and to the 
members of the Subcommittee for their unstinting 
efforts in reviewing the reports and their helpful 
comments. 

The Subcommittee is grateful to World Wildlife 
Fund Canada, the Canadian Wildlife Service, and 
the National Museums of Canada (now Canadian 
Museum of Nature), for their assistance in the 
process. A special mention to Francis Cook and The 



Canadian Field-Naturalist for assistance in 
publication and editing and to all members of 
COSEWIC for their dedication and interest in the 
future of Canada's fauna and flora. Last, but not 
least, we gratefully acknowledge the financial and 
secretarial support provided through the Depart- 
ment of Fisheries and Oceans and the financial 
contribution of Fisheries and Oceans, Environment 
Canada, and World Wildlife Fund Canada, which 
has permitted the production of several new reports. 

Literature Cited 

Campbell, R. R. 1984. Rare and endangered fishes of 
Canada: The Committee on the Status of Endangered 
Wildlife in Canada (COSEWIC) Fish and Marine 
Mammals Subcommittee. Canadian Field-Naturalist 
98(1): 71-74. 

Campbell, R. R. 1985. Rare and endangered fishes and 
marine mammals of Canada: COSEWIC Fish and 
Marine Mammals Subcommittee Reports: II. Canadian 
Field-Naturalist 99(3): 404-408. 

Campbell, R. R. 1987. Rare and endangered fishes and 
marine mammals of Canada: COSEWIC Fish and 
Marine Mammal Subcommittee Reports: III. Canadian 
Field-Naturalist 101(2): 165-170. 

Campbell, R. R. 1988. Rare and endangered fishes and 
marine mammals of Canada: COSEWIC Fish and 
Marine Mammal Subcommittee Reports: IV. Canadian 
Field-Naturalist 102(1): 81-86. 

Campbell, R. R. 1989. Rare and endangered fishes and 
marine mammals of Canada: COSEWIC Fish and 
Marine Mammals Subcommittee Reports: V. Canadian 
Field-Naturalist 103(2): 147-239. 

Cook, F. R. and D. Muir. 1984. The Committee on the 
Status of Endangered Wildlife in Canada (COSEWIC): 
History and progress. Canadian Field-Naturalist 98(1): 
63-70. 

Accepted 10 October 1989 



Status of the Fourhorn Sculpin, Myoxocephalus quadricornis, 
in Canada* 



J. Houston 

374 Fireside Drive, Woodlawn, Ontario KOA 3M0 

Houston, J. 1990. Status of the Fouriiorn Sculpin, Mvoxocephalus quadricornis, in Canada. Canadian Field- 
Naturalist 104(1): 7-13. 

The Fourhorn Sculpin, Myoxocephalus quadricornis, is a widespread marine species, closely related to, and the 
probable progenitor of, the freshwater Deepwater Sculpin, Myoxocephalus thompsoni. The species is widely 
distributed and abundant in the Canadian Arctic and is an important forage species for other fish and for waterfowl. 
Marine populations are secure but evolving isolates in euryhaline lakes of the Canadian Arctic Archipelago are 
probably vulnerable as these lakes are sensitive to perturbations. 

Le chabot a quatre cornes, Myoxocephalus quadricornis, est une espece marine qui s'apparente etroitement au Chabot 
de profondeur, Myoxocephalus thompsoni, dont il est probablement I'ancetre. Largement et abondamment repandu 
dans I'Arctique canadien, il constitue une source alimentaire importante pour la sauvagine et les autres especes de 
poisson. Ce poisson marin pelagique est stable, mais les populations vivant dans les lacs euryhalins de I'Archipel 
arctique canadien sont probablement vulnerables etant donnee que ces eaux sont sensibles aux perturbations. 

Key Words: Fourhorn Sculpin, Myoxocephalus quadricornis, chabot de quatre cornes, scorpion fish, cottidae, 
sculpins, rare and endangered fishes. 



The Fourhorn Sculpin, Myoxocephalus quadri- 
cornis (Linnaeus 1738), originally described from 
the Baltic Sea, is a marine sculpin with a 
circumpolar Palearctic and Nearctic distribution 
(McAllister 1980). The species is closely related to 
the freshwater Deepwater Sculpin, Myoxocephalus 
thompsoni (Girard 1852). Their taxonomy has been 
the subject of several taxonomic and zoogeographic 
studies (Berg 1949; Walters 1955; McAllister 1961; 
Johnson 1964; McAllister and Aniskowicz 1976). 
McAllister and co-workers (McAllister 1961; 
McAllister and Aniskowicz 1976; McAllister et al. 
1978) examined specimens of both forms and the 
various postglacial Arctic freshwater relicts which 
had been described (McAllister 1961; Hubbs and 
Lagler 1964; Johnson 1064; McPhail and Lindsey 
1970; Dadswell 1972) and considered marine and 
freshwater forms as distinct species based on 
morphological characteristics, distribution and 
ecology. The postglacial Arctic relicts fell within the 
definitions of the marine Fourhorn Sculpin. 

A status report has previously been completed on 
the freshwater form (Parker 1988). The present 
report deals solely with the "marine form" to which 
the name Myoxocephalus quadricornis is now 
restricted. The common and scientific names used 
here are those adopted by the American Fisheries 
Society (Robins et al. 1980). 

Description 

The Fourhorn Sculpin (Figure 1) is larger than 
the Deepwater Sculpin. It may attain a total length 



of up to 34.0 cm, but most specimens are 
somewhat smaller (McAllister 1980). The species 
can usually be easily distinguished from other 
cottids by the presence of four long protuberances 
(frontal and parietal spines) on the head (Scott and 
Scott 1988), which give rise to the name. There are 
also four well-developed preopercular spines as 
well as nasal and cleithral spines. The body is 
elongate with a slender caudal peduncle. The head 
is flattened and wide with close set eyes on the top 
of the head. There are two dorsal fins, the first 
being smaller and spiny; the caudal fin is square or 
truncate; the anal is soft rayed and has a long base; 
the pelvics are small, located well forward beneath 
the pectorals and have one spine and three to four 
soft rays; the pectorals are large and fanlike {see 
McAllister 1961; Scott and Scott 1988). 

These fish lack typical scales and may have 
tubercles, sometimes reduced to prickles, above 
and below the lateral line which seldom extend 
past the insertion of the second dorsal fin 
(McAllister 1961). The second dorsal fin is usually 
larger in mature males than in the female and the 
pelvics are also notably larger. There may be 
tubercles on the second dorsal and pectoral fins 
which are not found on females (McAllister 1961; 
McPhail and Lindsey 1970). 

The overall colouration is dark grey to brown, 
the back being darker, becoming lighter along the 
sides and light ventrally. The back and sides may 
be speckled or mottled and there are usually four to 
seven diffuse, saddle-like bands along the back and 



♦Vulnerable status approved and assigned by COSEWIC for those populations in landlocked lakes of the Arctic 
Archipelago II April 1989. 



The Canadian Field-Naturalist 



Vol. 104 




Figure \. Drawing of the Fourhorn Sculpin, Myoxocephalus quadricornis, kmale from 
Peterson Bay, King William Island, Northwest Territories (231 mm standard 
length) [drawing by C. Douglas, courtesy of D. E. McAllister, Canadian Museum 
of Nature]. 



sides. The pectoral fins may have up to three 
diffuse darker bands, the pelvics may be spotted, 
and the dorsal and anal fins blotched. The caudal 
fin usually shows dark brown mottling. Males 
develop a rosy colouration under the head, on the 
lower pectoral fin, and on the anal and pelvic fins 
(McAllister 1961; Scott and Scott 



Distribution 

The Fourhorn Sculpin has a circumpolar marine 
distribution from the Barents to the Bering Seas in 
the Palearctic and from Alaska to Labrador in the 
Nearctic. Although present in eastern Greenland, 
the species is absent from Iceland, Norway and 
Spitzbergen (McPhail and Lindsey 1970; McAllis- 
ter 1980). It occurs in both salt and brackish water, 
usually along the coast, but may be found up rivers 
as far as 150 km from the sea, and as landlocked 
relicts in some lakes (McAllister 1980). 

In Canada, the species has been reported from 
many points along the arctic coast from Alaska 
through to Labrador {see Figure 2; National 
Museum of Natural Sciences (now Canadian 
Museum of Nature (NMC); Royal Ontario 
Museum (ROM): Ontario Ministry of Natural 
Resources (OMNR), records) including Hudson 
and James bays and most islands of the Arctic 
Archipelago (see also Leim and Scott 1961; 
McAllister 1961, 1980; McPhail and Lindsey 1970; 
McAllister and Aniskowicz 1976). The species has 
been recorded some 200 km up the Mackenzie 
River and in streams on Victoria, Baker and 
Cornwallis islands and north to northern 
Ellesmere Island (McPhail and Lindsey 1970). 
Johnson (1964) reported the presence of A/yoxoce- 
phalus quadricornis in three lakes in Victoria 
Island and it has also been reported from lakes on 
Cornwallis Island and other islands in the 
archipelago (McAllister and Aniskowicz 1976). 

Protection 

No specific protective measures arc in place in 
either Canada or the United States. General 



protection is provided in Canadian waters under 
the Fisheries Act. 

Population Sizes and Trends 

Information on the size and trends of Fourhorn 
Sculpin populations is limited mostly to presence- 
absence records. Most sampling has not been 
extensive or sequential, but at sites which have 
been visited in each of the last three decades 
(Figure 2, NMC; OMNR; ROM). The species 
appeared to be consistently common at most 
locations. 

Craig and McCart (1976), in an overview offish 
utilization of nearshore habitats for the Beaufort 
Sea coasts from the Colville River (Alaska) to the 
Mackenzie River (Northwest Territories), found 
the Fourhorn Sculpin to be widely distributed. For 
some areas they suggested that it often was the only 
fish to be found and very abundant. Lawrence et al. 
(1984) found the species to be abundant at 11 of 23 
Tuktoyaktuk Peninsula locations where catches in 
1978, 1979, and 1980, accounted for 2.8 to 37.2% of 
the fish taken depending on season and type of 
gear. Fyke nets provided better catches per unit 
effort than gill nets, seines or plankton nets. In 
early spring and midsummer collections, mature 
females were dominant and the total fish caught 
was less than in midsummer when juveniles were 
abundant (Lawrence et al. 1984). Lawrence et al. 
( 1 984) found no other species as abundant with the 
exception of the Arctic Flounder {Liopsetta 
glacialis). Fall collections provided the highest 
catches and the greatest number of sexually mature 
fish. The species appeared to avoid the Mackenzie 
plume and was scarce in collections in the inner 
Kugmallit Bay, although small numbers did move 
I km or so into coastal streams, but not beyond the 
area of tidal influence (Lawrence et al. 1984). 
Ratynski (1983) did find larval Fourhorn Sculpin 
in Tuktoyaktuk Harbour. None were recorded 
outside the harbour in Kugmallit Bay. 



1990 



HOUSTON: Status of the Fourhorn Sculpin 




^~^~'\jX_ ^ '■ 'I --~-.^ I'^s c^-' i^' ^-. • J,'c; -C "'-fOi^. 



•fv 



-4G 






;:rv 




Figure 2. North American distribution of the Fourhorn Sculpin, Myoxocephalus quadricornis (from 
sources quoted in the text). O Alaska; • Pre 1960 collection records; A 1970's records; ■ Post 
1970 records. 



All in all, Fourhorn Sculpin are common in the 
Beaufort Sea (Ratynski 1983). Larvae and fish of 
all ages have been reported along the coasts of 
Alaska (Craig and McCart 1976; Craig etal. 1985), 
the Yukon coast (Kendel et al. 1975; Griffiths et al. 
1975), and along the Tuktoyaktuk Peninsula 
(Jones and Den Beste 1977; Ratynski 1983; 
Lawrence et al. 1984). Fourhorn Sculpin are 
known to overwinter in bays and inlets along the 
coast (Lawrence et al. 1984). 

Unfortunately, such detailed data are not 
available for the remainder of the Canadian Arctic 
coast, the eastern Arctic or the Arctic Archipelago. 
The Beaufort Sea has received considerable 
attention due to the Environmental Impact 
Assessment required by law in relation to 
hydrocarbon exploration and development in the 
area. The pace of activity has not been as great in 
the remainder of the Arctic and where sampling 
has been conducted, in Lancaster Sound and the 



Wellington Channel, the waters are deeper and the 
species not as Ukely to be found. Perhaps future 
activity in other areas of the Arctic will show the 
species to be as abundant there as in the west. 
Limited observations on or about Victoria, 
Cornwallis, Little Cornwallis and Ellesmere 
islands have indicated the presence of the species in 
these areas (McAlhster 1961). Welch (1985) 
observed that Fourhorn Sculpins were not taken in 
freshwater and were not known from the 
Saqvaqjvac area of Northern Hudson Bay, but 
were common in the high Arctic in lakes that 
contain anadromous Arctic Char {Salvelinus 
alpinis). 

Lake populations in the Arctic appear to be 
resident and to have been separated from their 
marine progenitors for several thousand years. 
While conspecific with the marine form, they do 
share distinctions and may be genetically 
differentiated. Their status should be evaluated 



10 



The Canadian Field-Naturalist 



Vol. 104 



separately from the marine form (D.E. McAllister, 
personal communication). 

Habitat 

The Fourhorn Sculpin is a marine species, usually 
found in salt and brackish waters close to the coast 
in waters from 20 m to the intertidal zone 
(McAllister 1980). McAllister (1980) reported that 
the species may run up estuaries as far as 150 km 
from the sea but do not form resident populations in 
fresh water. Lawrence et al. (1984) indicated that the 
species avoided the Mackenzie Plume and that they 
did not seem to move more than 1 km into coastal 
streams or beyond the region of tidal influence. 

Relict populations exist in some freshwater 
lakes or islands of the Arctic Archipelago, chiefly 
some lakes on Victoria, Cornwallis and EUesmere 
islands (Johnson 1964; McAllister and Aniskowicz 
1976) and perhaps others. Some of these lakes are 
meromictic, others are fresh, and McAllister and 
Aniskowicz (1976) considered that these were 
introduced by marine inundation following 
glaciation and isolated in freshening lakes during 
subsequent isostatic rebound. Occurrence of the 
species in these lakes is concurrent with that of the 
relict isopod Mesidosethra entomon glacialis 
which is not found in inland freshwaters on the 
mainland (McAllister and Aniskowicz 1976). 

These authors suggested that these relict 
populations arose independently and more 
recently from the Fourhorn Sculpin than did the 
Deepwater Sculpin and should still be referred to 
as Myoxocephalus quadricornis; although 
morphologically different, the variations are not 
sufficiently significant to warrant subspecific rank 
(McAllister and Aniskowicz 1976). Variation 
between such relict populations would be greater 
than in the sea, as there would be little, if any, gene 
flow between lake populations, allowing genetic 
differences to accumulate in each, in contrast to 
marine populations which have potential for 
continuous gene flow. Differences in body lengths 
and mean meristic values suggest genetic 
differences exist. Lake temperatures and salinity in 
meromictic lakes fluctuates, especially during egg 
development and thus influences variability in 
characters such as the number of vertebrae {see 
McAllister and Aniskowicz 1976). 

Adults of the species are benthic and the larvae 
are pelagic (Khan 1971; Khan and Faber 1973). 
During the open water season, juvenile and adult 
fish move into the nearshore brackish waters, seek 
preferred temperatures and salinity, and feed 
primarily on epibenthic invertebrates. Lawrence et 
al. (1984) reported that Fourhorn Sculpin caught 
from 1978 to I980along thc'l uktoyakluk I'cninsula 
in June and July were at locations where 
temperatures ranged from I to 9'^C and salinity was 



from 1 to 30. The largest catches occurred where 
salinity was above 10 and temperature less than 
6.5° C. During August 1980, salinities ranged from 
0.4 to 26.9 at capture sites and the largest numbers 
occurred where the salinity was 26.9 and the 
temperature 7.3° C (Lawrence etal. 1984). In the fall, 
these authors reported large catches at salinities of 3 
to 27 and temperatures from 0.4 to 6.0° C. Craig et 
al. (1985) reported similar movements to coastal 
areas in the open water season. 

The pelagic larvae have been reported to be 
common in the Beaufort along the coast {see 
Ratynski 1983) from Alaska to the Mackenzie 
Delta. They are often the most abundant larvae 
found in ichthyoplankton sampling (Jones and 
Den Beste 1977). Larvae have been taken along the 
Tuktoyaktuk Peninsula and in the Tuktoyaktuk 
harbour from July to September by plankton nets, 
seines and trawl {see Ratynski 1983). The larvae 
apparently appear early in the spring at the higher 
temperatures and lower salinities above the 
halocline (Ratynski 1983). 

Fourhorn Sculpins overwinter in bays, inlets 
and deltas along the coast in areas free of landfast 
ice (Lawrence et al. 1984; Craig et al. 1985). 
Overwintering fish have been taken in areas with 
salinities of 4.9 to 9.6 and temperatures less than 
0°C (Lawrence et al 1984; Craig et al. 1985). One 
female was taken in Tuktoyaktuk harbour in 
January 1981 at a depth of 9.5 m, salinity of 22.5, 
and temperature of less than 0°C (Lawrence et al. 
1984). While the species is known to overwinter in 
bays and inlets, the importance of offshore areas is 
not known. The habitat in the eastern Arctic has 
not been as well described, but one might assume 
that similar conditions prevail. 

General Biology 

McAllister(1980) reported amaximum age of 14 
years for the species. Based on calculations from 
the Tuktoyaktuk Peninsula, age at maturity would 
appear to be four to five years (Craig and McCart 
1976; Lawrence et al. 1984). No sexual differences 
in age at maturity were reported, but 70 to 88% of 
the midsummer catch were females in 1978 to 1980. 
Lawrence et al. ( 1 984) also noted that females grew 
significantly faster and lived longer (up to 14 years) 
than males (up to 1 1 years). From the data 
presented by Lawerence et al. ( 1984), length (total 
length) at maturity was 180 mm for females and 
about 150 mm for males. Maximum lengths of up 
to 340 mm have been reported (McAllister 1980) 
based on data from Alaska, the Yukon and Siberia 
(Andriyashev 1954; Percy 1975; Griffiths et al. 
1975, 1977). Similar information is not available 
from the eastern Arctic. 

The species spawning period and habitats are not 
known. The species probably spawns in late winter 



1990 



HOUSTON: Status of the Fourhorn Sculpin 



11 



under the ice (Percy 1975). A ripe and running 
female was taken in Tuktoyaktuk Harbour in 
January of 1981 at a depth of 9. 5 m, salinity 22.5 and 
temperature less than 0°C (Lawrence et al. 1984). 
The timing may vary with locale and may be related 
to variations in local winter temperature and 
salinity. 

The eggs hatch in May to June and free swimming 
larvae are generally abundant in shallow pelagic 
areas and along the coast by July (Khan and Faber 
1973; McAllister 1980; Ratynski 1983). Young-of- 
the-year in the Beaufort Sea grow from 19 mm in 
July to 38.6 m in September (Lawrence et al. 1984). 
Similar findings have been reported from Siberia, 
the Yukon and Alaska (Andriyashev 1954; Percy 
1975; Griffiths et al. 1975, 1977). Juveniles were 
50 mm in length by the following spring, up to 
120 mm at age two, and 150 mm plus by age three 
{see Lawrence et al. 1984). Growth appeared to be 
slower after maturity but continued through all age 
classes (Lawrence et al. 1984). 

Little information is available on species 
movements, but there are indications that Fourhorn 
Sculpin move to the coast from offshore areas in the 
spring and return offshore in the fall (Lawrence et al. 
1984). These authors also reported the avoidance of 
freshwater plumes by Fourhorn Sculpin, but noted 
a tendency for it to move into coastal streams in the 
region of tidal influence. Fourhorn Sculpins have 
also been reported up to 150 km upstream in the 
Mackenzie (McAllister 1980), but these are 
probably not resident populations. Such move- 
ments may be in relation to feeding or a response to 
preferred temperature and salinities. 

The larvae are undoubtedly planktivorous and 
smaller fish feed on amphipods and copepods (Craig 
and McCart 1976). The diet of large fish consists of 
isopods, amphipods, mysids and plant material; 
isopods and amphipods being most frequent 
(McAllister 1961). Lawrence et al. (1984) also found 
fish (Fourhorn Sculpin) and pelecypods in stomachs 
of fish taken from the western Arctic. Chironomid 
larvae, annelids, fish eggs and fish (smelt, Elginus 
novaga, and Threespined Stickleback, Gasterosteus 
aculeatus) have also been reported in stomach 
contents (McAllister 1961; Hansson et al. 1984). 
Fourhorn Sculpins of some of the freshwater and 
euryhaline lakes of the Archipelago islands are 
found in association with the isopod Mesidosethera 
entomon glacialis and that these lakes are usually 
devoid of freshwater fish (McAllister and 
Aniskowicz 1976). 

Limiting Factors 

The species may be susceptible to chronic trace 
contaminant exposure, pesticide exposure or to 
shifts in species composition. Hansson et al. (1984) 
have indicated that sublethal effects of pollutants 



may be responsible for a high frequency (54%) of 
spinal abnormalities in Fourhorn Sculpins in 
polluted areas of the Baltic Sea. However, it is not 
known if these effects are of ecological significance 
or if they influence the survival, reproduction or 
niche of the affected individuals. Such problems are 
as yet minimal in the Canadian range of the species, 
but may become more prevalent, if and when 
development, particularly petrochemical explora- 
tion and exploitation, intensifies. An exception is 
Garrow Lake, Cornwallis Island, Northwest 
Territories. This lake is meromictic and hypersaline. 
The lower, oxygen-free zone is populated by 
bacteria which are the forage base for the Crustacea 
of the upper zones which dive into the lower zone to 
feed. These Crustacea form the forage base of the 
lake's Fourhorn Sculpin population, the only fish in 
the lake. Mining waste from a nearby lead-zinc mine 
is being pumped into the lake to prevent pollution of 
the Arctic Ocean, and consequently destroying the 
bacteria which are the base of the food chain 
(Dickman 1988). 

The demise of the freshwater form, Myoxocepha- 
lus thompsoni, in Lake Ontario has been attributed 
to chronic pesticide and herbicide pollution or re- 
cruitment failure as a result of predation or competi- 
tion by Alewife, Alosa pseudoharengus (Christie 
1973; Gray 1979). The Fourhorn Sculpin is 
apparently subject to intense predation in shallow 
water by gulls, waterfowl and shorebirds. The species 
is also eaten by whitefish {Coregonus sp), Arctic 
Char, eelpouts and other Cottidae including other 
members of its own species {see McAllister 1961). 

Special Signiflcance of the Species 

The Fourhorn Sculpin is of little direct 
commercial or sportfishing interest, but it plays an 
important role in the shallow coastal Arctic 
ecosystem, where it is often the dominant species. It 
is of special interest in analysis of Canadian post- 
glacial dispersion patterns. It may be of value as an 
indicator of environmental quality due to the 
morphological changes that result from sublethal 
effects of pollutants and could be a key species to 
monitor in areas of development in the Arctic. 
Fourhorn Sculpins also provide a vital link in the 
Arctic marine environment in the conversion and 
transport of energy to higher trophic levels as it is an 
important component in the diet of waterfowl and 
fishes. Its distribution and evolution in the 
freshwater and euryhaline lakes of the Arctic Islands 
is of scientific interest and the populations should be 
protected in these particularly sensitive 
environments. 

Evaluation 

The Fourhorn Sculpin is common and abundant 
in Canadian Arctic coastal waters. More detailed 



12 



The Canadian Field-Naturalist 



Vol. 104 



information is required on distribution, abun- 
dance, biology and ecology, but the species is 
secure in Canada. 

Evolving populations in the freshwater and 
euryhahne lakes of the Arctic Archipelago are 
probably genetically unique and should be given 
special consideration. These populations are 
vulnerable. The population in Garrrow Lake, 
Cornwallis Island, is in danger of extirpation or 
may already have been lost. 

Acknowledgments 

The author gratefully acknowledges the 
Department of Fisheries and Oceans for their 
support in the production of this report. Thanks 
are also due Don McAllister, National Museum of 
Natural Sciences (now Canadian Museum of 
Nature), G. Gale, Ontario Ministry of Natural 
Resources, and E. Holm, Royal Ontario Museum 
for provision of information on collection records. 

Literature Cited 

Andriyashev, A. P. 1954. Ryby servernykh murei 
SSSR. [Fishes of the northern seas of the USSR] 
Fauna SSSR 53: 1-567. [Israel Program for Scientific 
Translations, Number 836. 1964]. 

Berg, L. L. 1949. Ryby presmykh vod SSSR i sopredel 
Tnykh stran. [Freshwater fishes of the USSR and 
adjacent countries, Volume III] Akademiya Nauk 
USSR Zoological Institute Guide to the Fauna of the 
USSR. Number 30. [Israel Program for Scientific 
Translations, Number 743. 1967]. 

Christie, W. 1973. A review of the changes in the fish 
species composition of Lake Ontario. Great Lakes 
Fishery Commission Technical Report 23: 65. 

Craig, P. C, and P. McCart. 1976. Fish use of 
nearshore coastal waters in the western Arctic: 
emphasis on anadromous species. Pages 361-388 in 
Assessment of the Arctic marine environment: selected 
topics. Edited by D. W. Hood and D. C. Burwell. 
Institute of Marine Science, University of Alaska, 
Fairbanks, Alaska, Occasional Paper Number 4. 

Craig, P. C, W. B. Griffiths, L. Haldorson, and H. 
McElderry. 1985. Distributional patterns of fishes in 
an Alaskan Arctic lagoon. Polar Biology 4: 9-18. 

Dadswell, M. J. 1972. Post-glacial dispersal of four 
freshwater fishes on the basis of new distribution 
records from eastern Ontario and western Quebec. 
Journal of the Fisheries Research Board of Canada 29: 
545-553. 

Dickman, M. 1988. Death of a lake. Globe and Mail, 7 
March 1988: Al, A3. 

Girard, G. 1 852. Contributions to the natural history of 
the freshwater fishes of north America. I: A 
monograph of the cottids. Smithsonian Contributions 
to Knowledge 3(article 3). 

Gray, J. E. 1979. Lake Ontario Tactical Fisheries Plan. 
Resource Document Number 9. Coldwater commun- 
ity rehabilitation: (I) Sea Lamprey, (2) Alewife, (3) 
Smelt, (4) Sculpins, (5) Deepwater Ciscos. Ontario 
Ministry of Natural Resources, Toronto, Ontario. 



Griffiths, W. P., P. Craig, G. Walder, and G. Mann. 

1 975. Fisheries investigations in a coastal region of the 
Beaufort Sea (Nunaluk Lagoon, Yukon Territory). 
Arctic Gas Biological Report Series 34 (2): 7-219. 

Griffiths, W., J. Den Beste, and J. Craig. 1977. Fisheries 
investigations in a coastal region of the Beaufort Sea 
(Kaktovik Lagoon, Barter Island, Alaska). Arctic Gas 
Biological Report Series 41: 1-116. 

Hansson, S., B-E. Bengtsson, and A. Bengttsson. 1984. 
Stomach contents in Baltic fourhorn sculpin 
{Myoxocephalus quadricornis L.) with normal and 
deformed spinal vertebrae. Marine Pollution Bulletin 
15(10): 315-317. 

Hubbs, C. L., and K. F. Lagler. 1964. Fishes of the 
Great Lakes region. University of Michigan Press, 
Ann Arbor, Michigan. 

Johnson, L. 1964. Marine glacial relicts of the 
Canadian arctic islands. Systematic Zoology 1 3(7): 76- 
91. 

Jones, M. L., and J. Den Beste. 1977. Tuft Point and 
adjacent coastal areas fisheries project. Unpublished 
Report for Imperial Oil Limited, Aquatic Environ- 
ments Limhed, Calgary, Alberta. 

Kendel, R. E., R. L. C. Johnston, U. Lobsiger, and 
M. D. Kozak. 1975. Fishes of the Yukon coast. 
Beaufort Sea Project Technical Report 6. 

Khan, N. Y. 1971. Comparative morphology and 
ecology of the pelagic larvae of nine Cottidae (Pisces) 
of the northwest Atlantic and St. Lawrence drainages. 
M. Sc. thesis, University of Ottawa, Ottawa, Ontario. 

Khan, N. Y., and D. J. Faber. 1973. A comparison of 
the deepwater and fourhorn sculpin, myoxocephalus 
quadricornis L. From North America. 1. Morphologi- 
cal development. Pages 703-712 in The early life 
history of fish. Edited by H.S. Baxter. Springer- 
Verlag, New York, New York. 

Lawrence, M. J., G. Lasko, and S. Davies. 1984. A 
survey of coastal fishes of the southeastern Beaufort 
Sea. Canadian Technical Report of Fisheries and 
Aquatic Sciences 1220. 

Linnaeus, C. 1758. Systema naturae per regna tria 
naturae, secundum classes, ordines, genera, species, 
cum characteribus, differentiis, synonymis, locis. 
Laurentii Salvii, Holmiae, 12th Edition, Volume 1. 

McAllister, D. E. 1961. The origin and status of the 
Deepwater Sculpin Myoxocephalus thompsonii, a 
Nearctic glacial relict. National Museum of Canada, 
Bulletin Number 1 72, Contributions to Zoology, 1959. 

McAllister, D. E. 1980. Myoxocephalus quadricornis 
(Linnaeus) Fourhorn Sculpin. Page 826 in Atlas of 
North American freshwater fishes. Edited by D. S. 
Lee, C. R. Gilbert, G. H. Hocutt, R. E. Jenkins, D. E. 
McAllister, and J. R. Stauffer, Jr. North Carolina 
State Museum of Natural History Biological Survey 
Publication 1980-12. 

McAllister, D. E., and J. Aniskowicz. 1976. Vertebral 
number in North American sculpins of the Myoxoce- 
phalus quadric(>rnis-cQmp\c\. Journal of the Fisheries 
Research Board of Canada 33: 2792-2799. 

McAllister, D. E., R. Murphy, and J. Morrison. 1978. 
The compleat minicomputer cataloging and research 
system for a museum. Curator 21(1); 63-91. 

McPhail, J. D., and C. C. Lindsey. 1970. Freshwater 
fishes of northwestern Canada and Alaska. Fisheries 
Research Board of Canada Bulletin 173. 



1990 



HOUSTON: Status of the Fourhorn Sculpin 



13 



Parker, B. 1988. Status of the Deepwater Sculpin, 
Myoxocephalus thompsoni, in Canada. Canadian Field- 
Naturalist 102(1): 126-131. 

Percy, R. 1975. Fishes of the outer Mackenzie Delta. 
Beaufort Sea Project Technical Report 8. 

Ratynski, R. A. 1983. Mid-summer ichthyoplankton 
populations of Tuktoyaktuk Harbour, N WT. Canadian 
Technical Report of Fisheries and Aquatic Sciences 
1218. 

Ricker, K. E. 1959. The origins of two glacial relict 
crustaceans in North America, as related to Pleistocene 
glaciation. Canadian Journal of Zoology 37(6): 871-893. 

Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker, 
E. A. Lachner, R. N. Lear, and B. Scott. 1980. A list of 
common and scientific names of fishes from the United 
States and Canada (Fourth edition). American Fisheries 
Society Special Publication Number 12. 



Scott, W. B., and E.J. Grossman. 1973. Freshwater 
fishes of Canada. Fisheries Research Board of Canada 
Bulletin 184. 

Scott, W. B., and M. G. Scott 1988. Atlantic fishes of 
Canada. Canadian Bulletin of Fisheries and Aquatic 
Sciences Number 219. 

Walters, V. 1955. Fishes of the western Arctic America 
and eastern Arctic Siberia. Taxonomy and Zoogeo- 
graphy Bulletin of the American Museum of Natural 
History 106 (article 5): 259-368. 

Welch, H. E. 1985. Introduction to limnological 
research at Saqvaqjvac, Northern Hudson Bay. 
Canadian Journal of Fisheries and Aquatic Sciences 
42 (8): 494-505. 



Accepted 10 October 1989 



Status of the Spoonhead Sculpin, Coitus ricei, in Canada* 



J. Houston 



374 Fireside Drive, Woodlawn, Ontario KOA 3M0 

Houston, J. 1990. Status of the Spoonhead Sculpin, Cottus ricei, in Canada. Canadian Field-Naturalist 104(1): 
14-19. 

The Spoonhead Sculpin, Cottus ricei, is the most distinct North American member of the genus and on this continent 
is most closely related to the European Cottus gobei s,^ec\QS group. The species is widespread east of the Great Divide 
ranging into Quebec and north to the mouth of the Mackenzie River and other drainages into Hudson and James bays. 
It has been recorded from all of the Great Lakes, but may be declining in Lake Erie and declining or extirpated from 
Lake Ontario. Elsewhere, the species is common where found and collection efforts in the 1970s have filled in many 
information gaps on the distribution in Ontario and Quebec. The distribution of the species appears to be closely 
related to glacial lakes created at the end of the Wisconsin period of glaciation. Post-glacial dispersion of the species 
may still be occurring as drainage systems adjust to isostatic rebound. 

Le chabot a tete plate, Cottus ricei, est le membre le plus distinctif du genre en Amerique du Nord et, sur ce continent, 
est fortement apparente au chabot de torrent, Cottus gobei, une groupe d'espece d'Europe. L'espece est largement 
repandue a Test de la ligne de partage des eaux ou elle atteint le Quebec et, au nord, I'embouchure du fleuve Mackenzie 
et d'autres bassins versants de la bale d'Hudson et de la baie James. Elle a ete signalee dans tons les Grands Lacs, mais 
pourrait bien etre en declin dans le lac Erie et en declin, ou disparue, dans le lac Ontario. Ailleurs, l'espece est courante 
dans les zones qu'elle habite et les collectes realisees au cours des annees 1970 ont permis de combler bon nombre de 
lacunes quant a sa repartition en Ontario et au Quebec. L'espece semble etroitement liee aux lacs glaciaires crees a la fin 
de la glaciation du Wisconsin. La dispersion postglaciaire de l'espece peut ne pas etre terminee, les bassins versants 
continuant de s'ajuster a I'equilibre isostatique. 

Key Words: Spoonhead Sculpin, Cottus ricei, chabot a tete plate. Rice's Sculpin, Cottidae, sculpins, status, Canada. 



The Spoonhead Sculpin, Cottus ricei (Nelson 
1876), is the most distinctive member of the genus 
in North America and is probably most closely 
related to the European Cottus gobei species group 
(McAllister and Lindsey 1961). The body (Figure 
1) is oddly shaped, with a flattened head and 
tubular trunk which is compressed laterally toward 
the tail. The caudal peduncle is narrow and the tail 
fin is slightly rounded and not forked. There are 
two dorsal fins, the first being smaller than the 
second, and a long anal fin. The pelvic fins are 
small and ventral while the pectorals are large and 
fanlike. The eyes are located on the top of the head 
and normal scales are lacking on the body which is 
more or less covered by small "prickles" which are 
sometimes restricted to a patch behind the pectoral 
fln. 

These are small fish, most specimens being 4 to 6 
cm in length but individuals of up to 13.4 cm have 
been recorded (Delisle and Van Vliet 1968; Scott 
and Grossman 1973). There appear to be clinal 
northeast variations in the number of vertebrae, fin 
rays, and in pigmentation (Scott and Grossman 
1973; McAllister and Parker 1980). In the lower 
Great Lakes, the general colouration is light brown 



to tan, but individuals further north are decidedly 
darker. There may be dark, saddle-like blotches 
dorsally, three along the base of the second dorsal 
fin and one on the caudal peduncle. The remainder 
of the back and sides are diversely speckled with 
brown spots which may also be found on the dorsal 
and caudal fins. The remaining fins are usually clear, 
but may exhibit various patterns of pigmentation in 
some watersheds {see Scott and Grossman 1973). 

The distribution and biology of many fishes in 
Ganada, especially deepwater forms, are not well 
known. This inhibits discussion of their zoogeo- 
graphy and has given rise to misconceptions 
concerning their status. Cottus ricei is undoubtedly 
one such species. The following is a summary of the 
available information and an informed statement of 
the current status of the species. 

Distribution 

The North American distribution of the 
Spoonhead Sculpin (Figure 2) is virtually restricted 
to Ganada. The species has been reported in all of 
the Great Lakes, including Lake Michigan, and 
from inland waters in the State of Michigan (Scott 
and Grossman 1973). 



*Rcccivcd and accepted by COSI-WK" I I April I9K9 no designation required. 



14 



1990 



HOUSTON: Status of the Spoonhead Sculpin 



15 




Figure 1. Drawing of the Spoonhead Sculpin, Cottus ricei [from Scott and Grossman 
(1973) by permission]. 



The Canadian distribution includes the St. 
Lawrence River and Great Lakes drainages, 
islands in James Bay, the Hudson Bay and Arctic 
drainage to the mouth of the Mackenzie River. In 
North America, the species is considered to be a 
glacial relict, surviving glaciation in an upper 
Mississippi refugium and perhaps in the upper 
Missouri as well (McPhail and Lindsey 1970; 
Dadswell 1972; McAUister and Parker 1980; 
Grossman and McAllister 1986). The species 
probably moved northward with the edge of the 
retreating ice and is now confined mainly to 
Ganada and the glacial lakes and rivers resulting 
from the retreat of the Wisconsin ice, dying out in 
the original parts of its range (McPhail and 
Lindsey 1970; Dadswell 1972). 

The known eastern limit of the range appears to 
be the St. Lawrence River in Quebec near the lie 
d'Orleans (Scott and Grossman 1973) and north to 
Lake Mistassini (Dadswell 1972). In Ontario, the 
Spoonhead Sculpin exists where there is suitable 
habitat, including the Great lakes, north to Lake 
Abitibi and the Severn River on Hudson Bay. It 
has been reported from brackish waters near 
Akimiski Island, James Bay (Keleher and 
Kooyman 1957; Ryder et al. 1964). The species has 
been reported in Lake Winnipeg and the 
Saskatchewan River in Manitoba and north to the 
Nelson River and Nueltin Lake. In Saskatchewan, 
Spoonhead Sculpins have been collected from the 
Saskatchewan River and Lakes Wollaston and 
Athabasca. It has a diverse distribution in Alberta 
where Spoonhead Sculpins have been taken from 
the Bow River and Waterton Lakes area in the 
southern part of the province. GoUections from the 
Milk River system are questionable and have not 
been verified (Scott and Grossman 1973). The 
species extends west to the continental divide and 
north to the Athabasca system and Lesser Slave 
Lake and the Peace River below the canyon. In 
British Columbia, it occurs in the northeast in the 
Fort Nelson and Maskwa Rivers and the Peace- 
Laird system, east of the continental divide. There 
is one questionable record from the Flathead River 
near the U.S. border [University of British 
Columbia: UBC 56-577], but its presence there was 



not noted by Peden and Hughes (1984). This 
record may be of the Shorthead Sculpin, Cottus 
confusus. Spoonhead Sculpins also occur in the 
Yukon Territory, in the Peel-Caribou systems of 
the Mackenzie River basin. 

In the Northwest Territories, the species has 
been found in the upper and lower Mackenzie 
River of the Arctic drainage and the Thelon River 
system of the Hudson Bay drainage (McAllister 
1962; Scott and Grossman 1973). 

Protection 

There are no specific protective measures for the 
Spoonhead Sculpin in Canada. General protection 
is given under the terms of the Federal Fisheries 
Act or various provincial wildlife legislation. The 
species is listed as of special concern in Montana 
and as a Protected Species in New York State 
(Johnson 1987). 

Population Sizes and Trends 

The distribution and biology of many fishes in 
Ganada are poorly known, especially where these 
are deepwater forms and/ or not of commercial 
interest. Due to certain habitat preferences or 
particular habits, some fishes like the Spoonhead 
Sculpin will not be caught when collecting with 
standard gear. Although widely distributed across 
Ganada, information on population sizes and 
trends for the Spoonhead Sculpin is generally 
limited to presence or absence data. 

In Ontario and Quebec, the species was 
originally reported from all of the Great Lakes 
(Delisle and Van Vliet 1968; Scott and Grossman 
1973) and, in a few widely scattered lakes 
suggesting a lacustrine habitat for this fish in 
southeastern Ganada. During the 1970s, a series of 
collections by Dadswell (1972) extended the 
known range of the species 700 km east to the 
northern end of Lake Mistassini, Quebec, and 
added 88 new locations for the species in Ontario 
and Quebec, closing many gaps between previous 
distribution records. 

In southeastern Ganada, the species is found 
more often in lakes than in rivers and appears to be 
predominately lacustrine. In western Ganada, the 



16 



The Canadian Field-Naturalist 



Vol. 104 



species is more commonly found in running water 
(McPhail and Lindsey 1970; Dadswell 1972). The 
apparent difference in habitat could possibly be an 
artifact of sampling or because there are few glacial 
and modern lakes in the west. Dadswell (1972), for 
example, did not examine areas of running water 
and most western collections have not concen- 
trated on lakes using otter trawls. It has been 
suggested that systematic surveys throughout 
Canada with adequate gear would possibly fill in 
more gaps in the known range. For example, the 
species has recently been reported from the Speed 
River in southwestern Ontario (Ontario Ministry 
of National Resources). The suggestion is, 
however, highly unlikely. The Speed River record 
could be a misidentification. This habitat 
difference is exhibited by other species such as 
Percopsis omniscomaycus and Hybopsis storeria- 
nus (Scott and Grossman 1973: 421, 681). 

Dadswell (1972) has discussed the influence of 
light. This fish occurs in the depths of clear lakes 
but at shallower levels in turbid lakes. In the 
northwest, lakes of glacial origin are not as 
plentiful as in the southeast and rivers tend to be 
cooler. 

Most collections [National Museum of Natural 
Sciences (now Canadian Museum of Nature 
(NMC), Royal Ontario Museum (ROM), Ontario 
Ministry of Natural Resources (OMNR)] have not 
shown the species to be abundant where taken. 
However, Dadswell (1972) found Spoonhead 
Sculpins to be moderately abundant in deep, 
stratified lakes and very abundant in the shallow, 
turbid lakes of the Ontario/ Quebec clay belt. No 
other abundance information is available and most 
collection records are not recent; many are prior to 
the 1960s. Where more recent information is 
available from the same watersheds the presence of 
the species has been reconfirmed and as indicated 
previously, many new sites have been added. 

Parker and McKee (1980) and D. E. McAllister, 
National Museum of Natural Sciences, Ottawa, 
Ontario; personal communication) indicate there 
may be some concern that the species has become 
rare or extirpated in Lake Ontario where it has not 
been recently collected. The species is given 
protected status in New York where it was known 
only from Lake Ontario (Johnson 1987). The 
Deepwater Sculpin (Myoxocephalus thompsoni) 
is now believed extirpated from Lake Ontario and 
has not been collected recently in Lake Erie (Scott 
and Crossman 1973; Gray 1979; Parker 1988). 
Spoonhead Sculpins in Lake Ontario may have 
been affected by the same factors leading to the 
demise of the Deepwater Sculpin. Various theories 
for the disappearance of the latter have been 
suggested including pesticide or herbicide 
pollution, and prcdation by, or competition with 



Alewife {Alosa pseudoharengus) {see Scott and 
Crossman 1973; Gray 1979). Regardless, this 
species, unlike the former, is widely distributed 
elsewhere in Ontario and low numbers or 
extirpation in Lake Ontario, if real, would not 
threaten its existence in Ontario or Canada. 
Reintroductions, either natural or artificial, would 
be possible, providing the causative factor(s) is 
corrected, as the species has been recorded from 
the Lake Ontario watershed (NMNS; OMNR; 
ROM). However, the species disappearance from 
the two lower Great Lakes should be of concern, 
should the same unknown factor(s) affect the other 
lakes. 

Although many site records are based on a very 
few specimens, often from Lake Trout (Salvelinus 
namaycush) or burbot {Lota) stomachs, concen- 
trated collection efforts have detailed a wider 
distribution for the species {e.g. Dadswell 1972). 
With the possible exception of Lake Ontario, and 
perhaps Lake Erie, the species could be more 
widely distributed than present records indicate 
and should be looked for in other areas within the 
known range where suitable habitat exists. 

Habitat 

As mentioned previously, site records led 
McPhail and Lindsey (1970) to postulate a riverine 
preference in western Canada and Dadswell (1972) 
to suggest a lacustrine habitat in southeastern 
Canada. However, this may be solely the result of 
collection methods and choice of sites. Many older 
records, for instance, are from Lake Trout or 
burbot stomach contents (Scott and Crossman 
1973), and Dadswell (1972) did not sample in 
running water, although others have sampled 
many rivers in Ontario with similar results 
(McAllister, personal communication). 

The species appears to be a glacial relict, the 
distribution being closely associated with the 
maximum extent of the Wisconsin glacial lakes 
and their outlet channels (Dadswell 1972). 
Dispersal some 17 000 to 6 000 years ago was by 
means of the post-glacial waters north and east 
from a Mississippian refugium, the species dying 
out in southern parts of the range (McPhail and 
Lindsey 1970). Spoonhead Sculpins have 
dispersed somewhat beyond the glacial lake- 
marine areas and may be still dispersing (Dadswell 
1972) as drainages adjust to isostatic rebound plus 
stream capture or headwater piracy. 

Spoonhead Sculpins may be tolerant of turbid 
and brackish waters (Scott and Crossman 1973; 
McAllister and Parker 1980) but are usually taken 
in small swift streams, rivers and lakes (Scott and 
Crossman 1973). Depth preferences are not clear 
but are probably intermediate between those of the 
Deepwater Sculpin {Myoxocephalus thompsoni) 



1990 



HOUSTON: Status of the Spoonhead Sculpin 



17 














-< 



^^"^fV^^Q,^ 



r— <J -J^^^ 



^^% 



\ 



<^i,-. 









A 




Figure 2. Distribution of the Spoonhead Sculpin, Cottus ricei, in Canada. 



and the Slimy Sculpin (Co/m5 cognatus) in the Great 
Lakes (Scott and Grossman 1973), but depth is 
probably mediated by temperature and light (= water 
clarity). Deason (1979) gave a range for Lake 
Michigan from shore to about 140 m which appears 
to be most realistic (Scott and Grossman 1973). 

Delisle and Van Vliet (1968) found Spoonhead 
Sculpins in June and July, on the bottom from 17 
to 27 m, in Henry Lake, Gatineau Gounty, Quebec, 
at temperatures of 6.0 ot 7.2° G. In Pemichangan 
Lake, Gatineau Gounty, 12 Spoonhead Sculpins 
were taken at a depth of 42.4 m by the same authors 
in August at a temperature of 4.5° G. Dadswell 
(1972) reported that in deep, stratified lakes, the 
species was found at depths ranging from 5 to 50 m 
with a temperature range of 4 to 8°G. In shallow 
turbid lakes (Dadswell 1972), Spoonhead Sculpins 
were abundant at depths of 5 to 10 m with 
temperatures up to 18°G. 



General Biology 

Very little is known concerning the biology of the 
species. Scott (1967) considered the Spoonhead 
Sculpin to be a spring spawner. However, Delisle 
and Van Vliet (1968) suggested spawning in late 
summer or autumn. Ovaries examined from 
Ontario specimens support the latter view (Scott 
and Grossman 1973). Larvae of the family 
(Gottidae) have been described by Heufelder (1982). 

These fish are relatively small; most specimens are 
in the range of 4 to 6 cm (Scott and Grossman 1973). 
Specimens larger than 7 cm have been reported and it 
may be that smaller individuals are immature (Scott 
and Grossman 1973). Specimens of up to 10.2 cm 
have been taken in Lake Erie. The largest recorded 
specimen, 13.4 cm, was taken in Penichangan Lake, 
(Quebec (Delisle and Van Vliet 1968). 

No information is available on species 
movements or diet. Scott and Grossman (1973) 



The Canadian Field-Naturalist 



Vol. 104 



assumed that planktonic crustaceans and aquatic 
insect larvae would be important food items as for 
the Deepwater Sculpin (McAllister 1961; McPhail 
and Lindsey 1970). 

The parasitic fauna of the species is not known 
but the Spoonhead Sculpin is known to be 
important in the diet of Lake Trout and burbot 
(Deason 1939; Rawson 1959) and perhaps 
whitefish (Deason 1939). 

Limiting Factors 

The historical limiting factors for the Spoon- 
head Sculpin would appear to be the availability of 
Wisconsin glacial lakes, their outlet channels and 
interconnecting waterways. The species may be 
susceptible to chronic trace contaminant exposure, 
or to shifts in species compositions in the 
deepwater community, as indicated for the 
Deepwater Sculpin (see Parker 1988), but this 
remains to be shown. Habitat changes and 
degradation due to silting, etc., may be limiting 
distribution in southwestern Ontario. 

Special Significance of the Species 

The species is of little direct interest to the 
commercial or sport fisheries, but indirectly it is of 
interest as it is a traditional natural food of Lake 
Trout and burbot (Scott and Grossman 1973). 

Spoonhead Sculpin are also of special interest to 
the scientific community since the distribution 
provides information on the geological or glacial 
history of the region and an indication of 
environmental conditions. 

Evaluation 

The Spoonhead Sculpin is widely dispersed 
throughout the country. With the possible 
exception of Lake Ontario, and perhaps Lake Erie, 
regional populations are common where found. 
Additional information is required to establish the 
situation in Lakes Ontario and Erie. 

Acknowledgments 

Funding in support of this report was provided 
through the Department of Fisheries and Oceans. 
The author gratefully acknowledges the assistance 
of the National Museum of Natural Sciences (now 
Canadian Museum of Nature), the Royal Ontario 
Museum and the Ontario Ministry of Natural 
Resources for provision of information on 
collection records. 

Literature Cited 

Christie, W. 1973. A review of the changes in fish 
species composition of Lake Ontario. Great Lakes 
Fisheries Commission Technical Report 2.3: 65. 



Crossman, E. J., and D. E. McAllister. 1986. 
Zoogeography of freshwater fishes of the Hudson Bay 
drainage, Ungava Bay and the American Archipelago. 
Pages 52-104 in Zoogeography of North American 
freshwater fishes. Edited by E. O. Wiley. John Wiley 
and Sons, Toronto, Ontario. 

Dadswell, M. J. 1972. Postglacial dispersal of four 
deepwater fishes on the basis of new distribution records 
in eastern Ontario and western Quebec. Journal of the 
Fisheries Research Board of Canada 29: 545-553. 

Deason, H. J. 1939. The distribution of cottid fishes in 
Lake Michigan. Paper of the Michigan Academy 
Science, Arts and Letters 24(2): 105-1 15. 

Delisle, C, and W. Van Vliet. 1968. First records of the 
Sculpins Myoxocephalus thompsonii and Cottus ricei 
from the Ottawa valley, southwestern Quebec. Journal 
of the Fisheries Research Board of Canada 25(1 2): 2733- 
2737. 

Gray, J. E. 1979. Lake Ontario Tactical Fisheries Plan. 
Resource Document Number 9. Coldwater community 
rehabilitation. (I) Sea Lamprey, (2) Alewife, (3) Smelt, 
(4) Sculpins, and (5) Deepwater Ciscos. Ontario 
Ministry of Natural Resources, Toronto, Ontario. 

Heufelder, G. 1982. Family Cottidae, sculpins. Pages 
656-676 in Identification of larval fishes of the Great 
Lakes basin with emphasis on the Lake Michigan 
drainage. Edited by N. A. Auer. Great Lakes Fisheries 
Commission Special Publication 82-3. 

Johnson, J. E. 1987. Protectedfishesof the United States 
and Canada. American Fisheries Society, Bethesda, 
Maryland. 

Keleher, J. J., and B. Kooyman. 1987. Supplement to 
Hinks, "The fishes of Manitoba". Manitoba Depart- 
ment of Mines and Natural Resources, Winnipeg, 
Manitoba: 104-117. 

McAllister, D. E. 1961. The origin and status of the 
deepwater sculpin, Myoxocephalus thompsoni, a nea- 
arctic glacial relict. Bulletin of the National Museums of 
Canada Contributions to Zoology (1959) 172: 44-65. 

McAllister, D. E. 1962. Fishes of the 1960 "Salvelinus" 
program from western Arctic Canada. Bulletin of the 
National Museums of Canada Contributions to 
Zoology 185: 17-39. 

McAllister, D. E., and C. C. Lindsey. 1961. 
Systematics of the freshwater sculpins {Cottus) of 
British Columbia. Bulletin of the National Museums of 
Canada Contributions to Zoology (1959) 172: 66-89. 

McAllister, D. E., and B. Parker. 1980. Cottus ricei 
(Nelson), Spoonhead Sculpin. Page 832 in Atlas of 
North American freshwater fishes. Edited by D. S. Lee, 
C. R. Gilbert, G. H. HocuU, R. E. Jenkins, D. E. 
McAllister, and J. R. Stauffer, Jr. North Carolina State 
Museum of Natural History Biological Survey 
Publication Number 1980-12. 

McPhail, J. D., and C. C. Lindsey. 1970. Freshwater 
fishes of northwestern Canada and Alaska. Fisheries 
Research Board of Canada Bulletin 173. 

Parker, B. 1988. Status of the Deepwater Sculpin, 
Myoxocephalus thompsoni, in Canada. Canadian 
Field Naturalist 102(1): 126 131. 

Parker, B., and P. McKee. 1980. Rare, threatened and 
endangered fishes in southern Ontario: Status reports. 
Reports to the Department of Supply and Services, 
Department of Fisheries and Oceans, and National 
Museum of Natural Sciences. Beak Consultants 
Limited, Mississauga, Ontario. 



1990 



HOUSTON: Status of the Spoonhead Sculpin 



19 



Peden, A. E., and G. W. Hughes. 1984. Status of the 
Shorthead Sculpin, Cottus confusus, in the Flathead 
River, British Columbia. Canadian Field-Naturalist 
98(1): 127-133. 

Rawson, D. S. 1959. Limnology and fisheries of Cree 
and Wollaston Lakes in northern Saskatchewan. 
Fisheries Branch, Saskatchewan Department of 
Natural Resources, Saskatoon, Saskatchewan. 
Fisheries Report Number 4. 

Ryder, R. A., W. B. Scott, and E. J. Grossman. 1964. 
Fishes of northern Ontario, north of the Albany River. 



Royal Ontario Museum Life Sciences Contributions 

Number 60. 
Scott, W. B. 1967. Freshwater fishes of eastern Canada. 

Second edition. University of Toronto Press, Toronto, 

Ontario. 
Scott, W. B., and E.J. Grossman. 1973. Freshwater 

fishes of Canada. Fisheries Research Board of Canada 

Bulletin 184. 



Accepted 10 October 1989 



Status of the Bering Wolffish, Anarhichas orientalis, in Canada* 

J. Houston' and D. E. McAllister2 

'374 Fireside Drive, Woodlawn, Ontario KOA 3M0 
^Canadian Museum of Nature, Ottawa, Ontario KIP 6P4 

Houston, J., and D. E. McAllister. 1990. Status of the Bering Wolffish, Anarhichas orientalis, in Canada. Canadian 
Field-Naturalist 104(1): 20-23. 

The Bering Wolffish, Anarhichas orientalis, has a spotty distribution in the northwest Pacific Ocean, and the Bering 
Sea. The species has been recorded at one location in the Canadian Arctic from Bathurst Inlet. Very little is known on 
the biology and ecology of these fish and its shallow water habitat which are unusual among the members of the 
Anarhichaidae. The species should be considered vulnerable in Canada. 

Le loup de Bering, Anarhichas orientaUs, presente une aire de distribution sporadique dans le nord-ouest du Pacifique 
et dans la mer de Bering. Cette espece a ete observee en un seul endroit de I'Arctique canadien, Finlet Bathurst. On 
connait tres peu de choses sur la biologic et I'ecologie de ce poisson qui est rare parmi les Anarhichadides dans leur 
habitat en eau peu profonde. L'espece doit etre consideree vulnerable au Canada. 

Key Words: Bering Wolffish, Anarhichas orientalis, Loup de Bering, Anarhichadidae, wolffishes, arctic marine, rare 
and endangered fishes, status, Canada. 



Wolffishes (Anarhichadidae) are large blenny- 
like fishes of the deep, cold waters of the 
continental shelves of the Pacific, Arctic and 
Atlantic oceans. They have a generally laterally 
compressed and elongated shape tapering to a 
slender caudal peduncle. The body is usually naked 
or with a few small cycloid scales, the lateral line 
may be absent or reduced and pelvic fins are 
absent. These fish have strong conical canine teeth 
and molars (McClane 1978; Nelson 1984). 

The Bering Wolffish, Anarhichas orientalis 
(Pallas 1814), [Figure 1] differs from the five other 
species of the genera in having 53 or more anal 
rays, deeper more rounded caudal fins, and is 
found only in the North Pacific and western Arctic 
oceans (McAllister, unpublished manuscript. 
Marine Fishes of the Canadian Arctic). The 
vomerine row of teeth is also larger than the 
palentine rows and extends beyond the latter, the 
upper lip is wrinkled and not papillate. 

Members of the species have a short blunt head 
and steep snout with large canine teeth pointing 
forward from the end of the jaws. The body is long 
and compressed with small cycloid scales scattered 
over the surface. The lateral line canal and pores 
are absent, but there are instead three rows of 
neuromasts, a short predorsal row, a dorsolateral 
row extending to the pectoral and a short ventral 
row (McAllister, unpublished). The dorsal fin is 
long and curves down to the short, rounded caudal 
fin. The anal fin is also long and narrowly united 
with the caudal fin at the base, the pectorals are 



broad and oval in shape {see Barsukov 1959; 
McAllister, unpublished). The species is known to 
reach lengths of up to 12 cm and weigh from 3 to 
15 kg (Barsukov 1959). Bering Wolffish are 
reported to be dark brown in colour and usually 
without dark spots or stripes (Andriyashev 1954). 
Young fish may have horizontal lines on the back 
and dark spots on the head (Andriyashev 1954; 
McAllister, unpublished). 

The species is known from only one region of 
western Arctic Canada and has not been recorded 
elsewhere in the Canadian Arctic. Not only is the 
distribution limited but the species is apparently 
not abundant (Smith 1977). The following report 
on the species was prepared for the Committee on 
the Status of Endangered Wildlife in Canada 
(COSEWIC) to outline the status of this species in 
Canada. 

Distribution 

The known distribution of the species is spotty; 
the range is the northwestern part of the Pacific, 
The Gulf of Tatary, the Sea of Okhotsk, the Bering 
Sea north to Norton Sound near the Bering Strait, 
and in Bathurst Inlet in the western Canadian 
Arctic (Figure 2). The species is not found along 
the Pacific coast of North America (Andriyashev 
1954; Barsukov 1959; McAllister, unpublished). 

Protection 

No specific legislation is in place in any part of 
the known range. In Canada, general protection is 



♦Vulnerable status approved and assigned by COSIWIC 1 I April I9H9. 



20 



1990 



Houston and McAllister: Status of the Bering Wolffish 



21 




Figure 1. Illustration of the Bering Wolffish, Anarhichas orientalis (drawing by C. 
Douglas, courtesy D. E. McAllister, Canadian Museum of Nature). 



potentially available under the Habitat Section of 
the Fisheries Act. 

Population Sizes and Trends 

Very little information is available on the 
abundance of this species except for presence/ 
absence data. Andriyashev (1954) and Barsukov 
(1959) indicate that the species is common in the 
northwestern Pacific and Bering Sea. However, it 
cannot be that abundant as it is considered of no 
economic interest, whereas other species of the 
genus are exploited by the commercial fisheries 
(Andriyashev 1954). 

There are only three records (Canadian Museum 
of Nature collections) for the species in Canada 




Figure 2. General distribution of Bering Wolffish 
populations. 



(Figure 3), although wolffish (either Anarhichas 
orientalis or Anarhichas denticulatus) are known 
to Inuit of the area as the old womanfish (Smith 
1977). The known distribution of the species is 
spotty and raises the question as to whether this 
represents a dependence on localized ecological 
conditions or whether normal collection gear are 
unsuccessful in capturing them (McAllister, 
unpubUshed). As they are not well known to the 
Inuit of the area it is more likely that the former is 
the case, as a species with these characteristics 
would have been known to, and reported by the 
Inuit. In addition, fairly intensive sampling 
programs have been undertaken by various 
agencies (Fisheries and Oceans, Canadian 
Museum of Nature, and various consultants in 
response to the requirements for environmental 
impact assessments in relation to oil and gas 
exploration and development), especially in the 
shallower waters of the western Canadian Arctic 
(Hunter et al. 1 984) and these have failed to turn up 
specimens elsewhere in the Canadian Arctic. 

Habitat 

Bering Wolffish are not a well-studied species. 
Very little is known of their habitat. The species is 
usually found in shallower waters than the other 
members of the genus and seems to prefer coastal 
areas with algae covered rocky bottoms. They 
apparently keep to the shallower coastal areas 
from spring to late fall, moving offshore when the 
coast is ice-bound (Andriyashev 1954). 

The species would appear to be benthophagic 
(Barsukov 1959), feeding on crabs and molluscs. 
Their habits may restrict their distribution to 
highly localized sites where suitable conditions are 
available i.e., shallower water, warmer tempera- 
tures. The species is likely not found in the eastern 



22 



The Canadian Field-Naturalist 



Vol. 104 




Figure 3. Canadian collection records for the Bering Wolffish. 



Arctic because the waters of the nearshore areas 
are generally deeper and colder. 

General Biology 

The biology of the species is virtually unknown. 
One large female (1.12m) with well-developed eggs 
was taken in late May from Avachinskaya Bay, 
Kamchatka (Andriyashev 1954). Barsukov (1959) 
reported egg diameters at 4.0 to 4.5 mm. The 
larvae are apparently pelagic and an individual of 
19 mm in length was once cast aboard a ship 
during a storm in May in the Bering Sea 
(Andriyashev 1954). Kobayashi (1961) described 
young fish of the species based on two specimens 
collected from the Okhotsk Sea in June of 1957 
and August 1958, respectively. 

Maturity may be reached at 15 to 17 cm, a fish of 
4 1 cm was aged at 4+ years, one at 70 cm at 8+ years 
and a 1 12 cm female at 17+ years (Barsukov 1959). 

The species is a bottom dweller and although the 
diet has not been well studied, crabs and mussels 
are known to be important food items (Andriya- 
shev 1954). 

Limiting Factors 

Not known, however habitat requirements may 
be very narrow and thus limit distribution of the 
species. 

Special Significance of the Species 

The species is currently of little or no economic 
importance. Smith (1977) indicates that the Bering 
Wolffish may be a forage species for arctic seals. 
Dunbar (1970) suggests that wolffish are excellent 
to eat and that the hide could be tanned for 
commercial sale. However, the two Arctic species, 
Anarhichas orien talis and Anarhichas denticula- 
tus have a very narrow range and arc probably not 



abundant (Leim and Scott 1966; Dunbar 1970). 
Dunbar (1970) has also pointed out that the fishing 
potential of arctic waters is negligible so even if 
further populations are found in the Arctic there is 
little likelihood that the species would ever be 
found in commercial abundance. 

Evaluation 

Very little is known of this species in Canada or 
elsewhere. The Canadian distribution is limited 
and the species is probably not common in Arctic 
waters. Bering Wolffish are a naturally rare species 
in Canadian waters. The species should be 
considered vulnerable due to its narrow distribu- 
tion in the western Canadian arctic. 

Acknowledgments 

Financial support for the preparation of this 
report was provided by World Wildlife Fund 
(Canada), the Department of Fisheries and Oceans 
and Environment Canada. 

Literature Cited 

Andriyashev, A. P. 1954. Ryby sevemykh morei SSSR. 
Akademiya Nauk Soyuza Sovetskikh sotsialistiches- 
kikh Respublik, Moskoa Leningrad. [Fishes of the 
northern sea of the U.S.S.R.] Translated from 
Russian. Israel Program for Scientific Translations, 
.lerusalem 1964. 

Barsukov, V. V. 1959. Sem Zubalok (Anarhichadidae) 
Fauna SSR Akademiia Nauk USSR Zologicheskii 
Inslitut 73: 1-171. 

Dunbar, M. J. 1970. On the fishery potential of the sea 
waters of the Canadian north. Arctic 23(3): 150-174. 

Hunter, J. C, S. T. I,each, D. E. McAllister, and M. B. 
Steigerwald. 1984. A distributional atlas of records of 
the sea marine fishes of Arctic Canada in the National 
Museums of Canada and Arctic Biological Station. 
Syllogeus (National Museum of Natural Sciences) 52. 



1990 



Houston and McAllister: Status of the Bering Wolffish 



23 



Kobayashi, K. 1961. Youngof the wolf-fish /IwarA/c/ia^ 

orientalis Pallas. Bulletin of the Faculty of Fisheries, 

Hokkaido University 12(1): 1-4. 
Leim, A.M., and W. B. Scott. 1966. Fishes of the 

Atlantic coast of Canada. Fisheries Research Board of 

Canada Bulletin Number 155. 
McClane, A.J. Editor. 1978. Field guide to saltwater 

fishes of North America. Holt, Rinehart, Winston, 

New York, New York. 



Nelson, J. S. 1984. Fishes of the world. Second edition. 
John Wiley and Sons, Toronto, Ontario. 

Smith, T. G. 1977. The Wolffish, cf. Anarhichas 
denticulatus , new to the Amundsen Gulf area, 
Northwest Territories, and probable prey of the 
Ringed Seal. Canadian Field-Naturalist 91(3): 288. 

Accepted 10 October 1989 



Status of the Blackline Prickleback, Acantholumpenus mackayi, 
in Canada* 

J. Houston' and D. E. McAllister2 

'374 Fireside Drive, Woodlawn, Ontario KOA 3M0 

^Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario KIP 6P4 

Houston, J., and D. E. McAllister. 1990. Status of the Blackline Prickleback, Acantholumpenus mackayi, in 
Canada. Canadian Field-Naturalist 104(1): 24-28. 

The Blackline Prickleback, Acantholumpenus mackayi, is one of the larger eelblennies of the North Pacific and Arctic 
oceans. The Canadian range probably represents the northeastern fringe of the range and is limited to the Beaufort Sea 
in the vicinity of the Tuktoyaktuk Peninsula. The species is probably euryhaline, benthophagic and found over sand, 
mud or silty bottoms at depths to 60 m. Very little is known of the biology of the species, especially in Canadian waters 
where it is rare and vulnerable. The future of these fish in Canada is threatened by possible habitat loss or degradation 
through activities connected with the exploration and development of hydrocarbon deposits in the area. 

Le terrassier a six lignes, Acantholumpenus mackayi, est Tune des plus grosses lompenies de Tocean Pacifique Nord et 
de I'ocean Arctique. L'aire de repartition canadienne se situe probablement a la bordure est de I'aire de repartition 
globale et est limitee par la mer de Beaufort dans les environs de la peninsule de Tuktoyaktuk. II s'agit probablement 
d'une espece euryhaline, benthophage qu'on trouve sur des fonds sableux, vaseux ou limoneux a des profondeurs de 
60 m. Sa biologic reste tres mal connue, notamment dans les eaux canadiennes ou Tespece est rare et vulnerable. 
L'avenir de ce poisson au Canada est menace par la perte ou la degradation possible de son habitat a cause des activites 
associees a I'exploration et a Fexploitation des depots d'hydrocarbures dans le secteur. 

Key Words: Blackline Prickleback, Acantholumpenus mackayi, terrassier a six lignes, Spiny Eelblenny, Stichaeidae, 
lumpeniids, eelblennies, arctic marine fishes. 



The Blackline Prickleback, also known as the 
Spiny Eelblenny, Acantholumpenus mackayi 
(Gilbert 1893), is the only member of this genus of 
the prickleback family (Stichaeidae). Stichaeids 
are benthic fishes of the North Pacific, although a 
few genera are found in the Arctic and North 
Atlantic. The pricklebacks are not of importance 
in commercial or sport fisheries. They are usually 
rare, inhabiting deeper waters to depths of 1800 m 
or more, although some species may be found in 
intertidal areas (McClane 1978). Many of the 
species are small, but the Blackline Prickleback 
(Figure 1) may reach total lengths of 50 to 70 cm 
(McAllister, unpublished manuscript: The Marine 
Fishes of Arctic Canada). Acantholumpenus 
mackayi occurs in the Arctic and North Pacific 
oceans. The species was originally described by 
Gilbert (1895) as Lumpenus mackayi from 
specimens taken near Nushagak, Alaska and 
named for C. L. MacKay, a promising young, 
American ichthyologist who drowned at 
Nushagak in Bristol Bay in 1883 (McAllister, 
unpublished). 

The species differs from other members of the 
prickleback family in its pointed snout which 
projects beyond the upper lip. In addition, there 
are a greater number of dorsal spines (69 or more). 



There is a solid dark line along the base of the 
dorsal fin which does not exist in other members of 
the family (McAllister, unpublished). The body is 
long and slender and the dorsal fin extends almost 
the entire length of the body narrowly uniting with 
the caudal fin. The pelvic fins are thoracic, slender 
and short with one sharp spine; the pectorals are 
also well forward, but larger and oval shaped. The 
anal fin is also long and extends to the caudal fin 
with which it narrowly unites. 

Blackline Pricklebacks are yellowish to 
yellowish-brown with dark markings on the head 
and back. There is a more-or-less continuous dark 
band at the base of the dorsal fin and dark spots 
and lines on both sides of the lateral line; these 
disappear on the posterior third of the body. The 
fins are clear except for the caudal fin which is dark 
with a light posterior border (Gilbert 1895; Abe 
1972; McAllister, unpublished). 

Distribution 

The known distribution of the species is broken 
into three disjunct regions, each separated by over 
1 600 km of open ocean ( Figure 2). One area of the 
range is from the Sea of Japan to the Okhotsk Sea, 
another is the southern and eastern Bering Sea and 
the third is the Canadian Arctic (Legendre et al. 



^Vulnerable status approved and assigned by COSIWIC I 1 April I9S9. 



24 



1990 Houston and McAllister: Status of the Blackline Prickleback 



25 




......r^rrrrmmmji^^ 






asi^^fe.-^4i>ts^^^^ 





Figure \. Blackline Prickleback, Acantholumpenus mackayi, NMC65-0341-C, lateral, 
frontal and ventral views (drawing by C. Douglas, courtesy D. E. McAllister, 
Canadian Museum of Nature). 



1975) in the Beaufort Sea (Figure 3), and 
Mackenzie Delta in the area of Tuktoyaktuk 
(Lindberg and Krosyukova 1975; Shchetinnikov 
1983; McAllister, unpublished). Surveys to locate 
the species in eastern Kamchatka, the western 
Bering Sea, northern Alaska, the Yukon and 
elsewhere in the Northwest Territories have not 
produced evidence of intervening populations 
(McAllister, unpublished). Canadian populations 
probably represent the northeastern end of the 
species range. 

Protection 

No specific legislation exists for the protection 
of these fish in any part of the range. In Canada, 
general potential protection is available through 
the Habitat Section of the Fisheries Act. 

Population Sizes and Trends 

Very little information other than presence or 
absence data is available for this species. In 
Russian waters, the species has been reported in 
small numbers (Makuskov 1971; Shchetinnikov 
1983). Schetinnikov (1983) reported that the 
Blackline Prickleback is common in near-bottom 
waters of the Gulf of Terpenium around Sakhalin 
Island. Maximum population density was 50 to 
720 fish per km^ at depths of 20 to 50 m in the 
northern and central areas of the Gulf, but became 
less abundant towards the southeast and toward 
the coast (Shchetinnikov 1983). 

Similar information from Alaska and Canada is 
not available. Canadian Museum of Nature 
(NMS) collection records for Canadian specimens 
suggest that the species is not abundant in 
Canadian waters. Galbraith (1975) reporting on 
fish population surveys of the coastal Beaufort 
around Tuktoyaktuk Peninsula reported only two 



lumpenins from bottom otter trawls and less than 
100 from mid-water trawls. Ratynski (1983) 
reported only eight larvae from a series of trawls in 
1982 (Blackline Prickleback and Slender Eel- 
blenny, Lumpenus fabricii). These collections are 
the first records of lumpenin larvae in the Beaufort 
Sea and none were taken outside the harbour in 
Kugmallit Bay (Ratnyski 1983). Ratynski (1983) 
suggested that at the time of year of samphng, 
lumpenin samples may have been low as the larvae 
may have been large enough to avoid the nets or 
had moved from the area. 

Habitat 

Canadian specimens have been collected on mud 
bottoms at an average depth of 13 m (McAllister, 




Figure 2. General distribution of the Blackline 
Prickleback, Acantholumpenus mackayi. 



26 



The Canadian Field-Naturalist 



Vol. 104 




FlGl'RE 3. Canadian distribution of the Blackline Prickleback. Acatiiholunipemis niackayi. 



unpublisheif). In Alaska, it has been taken at the 
mouth of the Nushagak River to a depth of 56 m 
(Schmidt W50). The species is most abundant in 
the Gulf of Terpenium at 29 to 50 m over sandy 
and silty bottoms; it has also been taken from 
brackish waters in lakes of northern Hokkaido and 
the estuary of the Amur River (McAllister. 
unpublisht\D. The species thus appears to be 
tolerant of brackish waters but is likely euryhaline 
as salinities in the Gulf of Terpenium and the 
Aleutians would be expected to be high 
(McAllister, unpuhlisheif). Larval specimens were 
taken in Tukto\aktuk Harbour in the upper 
halocline at depths of 4.0 to 7.5 m (Ratynski 19S3) 
m cooler more saline waters. 

Shcheiinniko\ (1983) suggested that spawning 
areas arc located in shallower waters. Rat\nski 
(1983) indicated that the cooler saline bottom 
la>ers of Tuktovaktuk Harbour may be of critical 
importance for spawning of the Canadian 
populations. 



The body shape, frequency of catch in bottom 
trawling, and food composition all indicate that 
the Blackline Prickleback is benthic. feeding on the 
bottom and rarely in the water column near the 
bottom (Shchetinnikov 1983). Shchetinnikov 
(1983) also indicated that the species is non- 
schooling. 

General Biology 

Information on the biology of the species is 
limited. Ratynski (1983) reported that females 
taken in Tuktoyaktuk Harbour in September 1982 
w ere nearly ripe and females collected in .August of 
1971 had eggs 1.0 to 1.4 mm in diameter 
(McAllister 1975). Shchetinnikov (1983) indicated 
that males and females collected in July of 1978 in 
the Gulf of Terpenium were in post-spawning 
condition. Both Ratynski (1983) and Shchetin- 
nikov (1983) suggested that the fish move into 
shallow water to spawn and the younger fish 
remain in deepver waters to teed. 



1990 



Houston and McAllister: Status of the Blackline Prickleback 



27 



In the Gulf of Terpenium, males and females 
mature at a length of 30 to 40 cm and maximum 
length was found to be 62 cm (Shchetinnikov 
1983). McAllister {unpublished) suggested that 
males tend to be larger than females but 
Shchetinnikov (1983) found the reverse to be the 
case in the Gulf of Terpenium. In Canadian waters, 
the largest individual taken weighed 109 g and 
measured 49.4 cm (McAllister, unpublished). In 
Soviet waters, they may grow to 58.5 cm and up to 
70 cm off Japan (Ueno 1971). Based on the 
relatively low number of fish recorded it is not 
possible to make inferences as to taxonomic 
differences, but it does appear that fish from Japan 
may be larger, have more vertebrae, dorsal spines 
and rays and could represent a species or sub- 
species distinct from Acantholumpenus mackayi 
(McAllister, unpublished). 

Larval stages and growth have been outlined by 
Tokuya and Amaoka (1980). Larval lumpenids 
captured in July of 1982 at Tuktoyaktuk ranged in 
length from 15.8 to 21.5 mm (Ratynski 1983). 
Tokuya and Amaoka (1980) reported juvenile 
lengths at 36.0 to 38.5 mm in June off Hokhaido, 
Japan. 

Blackline Pricklebacks are typical bentho- 
phages, but in spite of their large size they have a 
small mouth and prey on smaller organisms. 
McAllister {unpublished) found that specimens 
taken in Canadian waters had been feeding on 
small clams 2 to 7 mm in length. Shchetinnikov 
(1983) reported that the species is not selective but 
feeds on all benthos of suitable size. The principal 
food items are sedentary and of limited mobility, 
mainly polychetes, gammarids and bi-valve 
molluscs. Camaceous gastropods and nudibranch 
molluscs are also included in the diet as are 
euphausids and young sea urchins (Shchetinnikov 
1983). Their specialized feeding on smaller prey 
permits the species to avoid competition with more 
active benthophages such as flounders (Shchetin- 
nikov 1983). 

Limiting Factors 

The species is of no economic importance and is 
not subject to exploitation except in Japan where 
they are used to make fish cakes and pastes 
(McAllister, unpublished). No other limiting 
factors are known for the population. However, 
perturbations in its near shore environment could 
have deleterious effects on spawning habitat and 
nursery areas of the larvae. In Canada, the species 
is not abundant and has been recorded only from 
the Mackenzie Delta in the region of Tuktoyaktuk 
Harbour which is also the centre of hydrocarbon 
exploration and development in the western 
Canadian Arctic. Dredging and drilling activities 
and vessel traffic in that harbour could very well be 



threatening to the continued existence of the 
species in Canadian water. 

Special Signiflcance of the Species 

The Blackline Prickleback is the only species in 
the genus. These fish are found only in the North 
Pacific and western Arctic oceans. The species is 
larger than many of the other lumpenids and lives in 
shallower waters. It is of no economic importance in 
Canada but is of interest due to its presence here, 
probably at the eastern fringe of its range. The three 
populations are disjunct and separated by over 
1600 km of open ocean. There is some indication 
that the most southwestern population in the Sea of 
Japan may be a distinct species or sub-species 
(McAUister, unpublished). If so, this would make 
the Canadian populations more unique. A detailed 
taxonomic study is required as perhaps all three 
populations are distinct. 

Evaluation 

The following factors are important in 
evaluating the status of this species in Canadian 
waters: (1) It has only been recorded, in Canada, in 
the Beaufort Sea in the area of the Mackenzie 
Delta; (2) The three populations of the species are 
disjunct, separated by some 1600 km of open seas. 
Each may be distinct; (3) The Canadian population 
probably represents the northeastern end of the 
range; (4) Except for the population of the Sea of 
Japan, these fish are not economically important; 
(5) The known Canadian range is in an area where 
the habitat is threatened by exploitation and 
development of hydrocarbon deposits. 

Although there are indications of reproductive 
success in the Canadian populations, numbers 
appear to be low, and based on this and the above, 
the species is deemed to be extremely rare and 
therefore vulnerable in Canada. 

Acknowledgments 

Financial support was provided by World 
Wildlife Fund Canada in cooperation with the 
Department of Fisheries and Oceans and 
Environment Canada. Thanks are also due to 
COSEWIC for the opportunity to present this 
material for their consideration. 

Literature Cited 

Abe, T. 1972. Key to the Japanese fishes. Revised third 
edition. Hokuryukam, Tokyo. 

Galbraith, D. 1975. Movements, distribution, popula- 
tions and food habits of fish in the coastal Beaufort 
Sea. Interim report of Project BI (Eastern). Arctic 
Biological Station, Ste. Anne de Bellevue, Quebec. 

Gilbert, C. H. 1895. The ichthyological collections of 
the steamer ''Albatross" during the years 1890 and 
1891. Reports of the U.S. Fisheries Commission (1 893) 
19: 392-476. 



28 



The Canadian Field-Naturalist 



Vol. 104 



Legendre, V., J. G., Hunter, and D. E. McAllister. 1975. 
French, English and scientific names of marine fishes of 
Arctic Canada. Syllogeus 7. 

Lindberg, G. U., and Z. V. Krasyukova. 1975. Fishes of 
the Sea of Japan and adjoining parts of the Okhotsk 
and Yellow Seas. [In Russian]. Nauka Press, 
Leningrad, Part 4: 423. 

Makuskov, M. E. 1971. Family Stichaeidae. Pages 514- 
516 in Animal Life, Volume 4. [In Russian]. 
Prosveshchenie Press, Moscow. 

Makuskov, V. M. 1958. The morphology and classifi- 
cation of the northern Blennioid fishes (Stichaeoidae, 
Blennioidei, Pisces). Proceedings of the Zoological 
Institute (Trudy Zoological Institute — Akademiya 
Nauk Soyuza Sovetskikh Satsialisticheskikh Respub- 
lik) 25: 3-129. [Translated from the Russian by the 
Ichthylogical Library, U.S. National Museum, 1959]. 

McClane, A. J. Editor 1978. McClane's field guide to 
saltwater fishes of North America. Holt, Rinehart and 
Winston, New York, New York. 



Ratynski, R. A. 1983. Mid-summer ichthyoplankton 
populations of Tuktoyaktuk Harbour, N.W.T. 
Canadian Technical Report of Fisheries and Aquatic 
Sciences 1218. 

Shchetinnikov, A. E. 1983. Nutrition of Acantholum- 
penus mackayi (Stichaeidae) in the Gulf of Terpenuim 
(Sakhalin Island). Journal of Ichthyology 6: 15-158. 

Tokuya, K., and K. Amaoka. 1980. Studies on larval 
and juvenile blennies in the coastal waters of the 
southern Hokhaido (Pisces; Blennioidei). Bulletin of 
the Faculty of Fisheries, Hokhaido University 31(1): 
16-49. 

Ueno, T. 1971. List of marine fishes from the waters of 
Hokhiado and its adjacent islands. Scientific Reports 
of the Hokhaido Fisheries Research Station 13: 
61-102. 

Accepted 10 October 1989 



Status of the Margined Madtom, Noturus insignis, in Canada* 
Cheryl D. Goodchild 

2168 Harcourt Crescent, Mississauga, Ontario L4Y IWl 

Goodchild, Cheryl D. 1990. Status of the Margined Madtom, Noturus insignis, in Canada. Canadian Field- 
Naturalist 104(1): 29-35. 

The Margined Madtom, Noturus insignis, is classified as threatened in Canada. In the early 1970s, it was first reported 
from isolated areas of the Ottawa River drainage in Ontario and Quebec, disjunct from the extreme northern fringe of 
its range in the United States. Its occurrence in Canada may be the result of accidental introduction of baitfish by 
anglers or of dispersal from New York State through the St. Lawrence River watershed. The population may be 
declining as attempts to recapture Margined Madtom in the 1980s were largely unsuccessful despite exhaustive 
sampling. Low numbers, fluctuations in spawning success and extremely specialized habitat requirements make this 
species' continued existence in Canada particularly perilous. 

Le chat-fou lisere, Noturus insignis, doit etre classe parmi les especes menacees au Canada. Au debut des annees 1970, 
ce poisson a ete pour la premiere fois signale dans des zones isole, du bassin versant de la riviere des Outaouais en 
Ontario et au Quebec, ce qui marque done une discontinuite par rapport a la limite la plus septentrionale de son aire de 
repartition aux Etats-Unis. Cette espece a peut-etre ete introduite accidentellement au Canada comme poisson d'appat 
utilise par les pecheurs sportifs ou bien elle s'est dispersee a partir de I'Etat de New York a travers le bassin versant du 
St-Laurent. La population de cette espece est peut-etre en train de decliner puisque, dans les annees 1980, on n'a pas 
bien reussi a recapturer le chat-fou lisere malgre les travaux d'echantillonnage exhaustifs. Le petit nombre de ces 
poissons, les fluctuations dans la reussite de la ponte et les conditions tres speciales qu'exige leur habitat rendent cette 
espece particulierement vulnerable. 

Key Words: Margined Madtom, Noturus insignis, chat-fou-lisere, catfishes, Ictaluridae, madtoms, rare and 
endangered fishes. 



The Margined Madtom, Noturus insignis 
(Richardson 1836), is a small ictalurid first 
reported from Canada by Rubec and Coad (1974). 
Although common in the upland Atlantic drainage 
of the United States, it has a very restricted 
distribution in Canada and occurs only in a few 
streams in the Ottawa River drainage. 

These fish (Figure 1) are slate-gray or yellowish 
shading to pale cream on the lower head and belly. 
There is a bridge of pigmented skin across the belly 
in front of the pelvic fins and the chin is dark in 
front of the barbels (Smith 1985). The margins of 
pectoral, anal, dorsal and caudal fins frequently 
have a dark band. The body is elongate, caudal 
peduncle deep, and head depressed. They are also 
distinguished by short teeth on posterior edges 
only of pectoral spines; lower jaw included; square 
caudal fin. Adults commonly attain 100 mm 
standard length (SL) with a specimen of 126 mm 
SL recorded from Virginia (Taylor 1969). Keys to 
identify the species in Canada are found in Rubec 
and Coad (1974) and Coad (1986). 

Distribution 

North America: The zoogeography and 
distribution of the species in the genus Noturus was 



discussed comprehensively by Taylor (1969). The 
greatest number of species of Noturus occur in 
upland regions of the east central United States. 
Because of favourable habitat, a band from 
Arkansas through Kentucky and Tennessee to 
Virginia and North Carolina was probably a refuge 
during Pleistocene glaciation for several northern 
species of Noturus. This may represent the center 
of their evolution and it appears most species were 
not materially affected by glaciation. Noturus 
insignis, an upland species, was probably restricted 
southward by the Piedmont Plateau in Georgia 
and South Carolina, but has successfully moved 
northward again. 

The native range of Noturus insignis includes 
part of the Lake Ontario drainage, from the Finger 
Lakes region southward through the Appalachian 
Highlands (Figure 2). This includes most of the 
Atlantic coastal streams from New York to 
Georgia, where they are most frequently found at 
or above the Fall Linef. Historical evidence points 
to the absence of N. insignis from the Upper Ohio 
and Tennessee systems (Figure 2). 

In recent years, Noturus insignis has been 
collected from the upper Ohio River system and 
from the Tennessee system where it is well 



♦Threatened status approved and assigned by COSEWIC 1 1 April 1989. 

fThe Fall Line is the line joining waterfalls on a number of approximately parallel rivers. In the eastern United States, 

it refers to the line running along a sharp increase of slope from the Atlantic Coastal Plain to the Appalachian 

Mountains. 



29 



30 



The Canadian Field-Naturalist 



Vol. 104 




Figure L Drawing of Margined Madtom, Noturus insignis (drawing by C. Douglas, 
courtesy D. E. McAllister, Canadian Museum of Nature). 



established. It is known from several localities in 
the Watauga drainage in North Carolina and from 
the North Fork of the Holston River, Virginia. It 
has been collected from three locaHties in the 
Monongahela and Youghiogheny system in West 
Virginia, Maryland, and Pennsylvania. Hocutt et 
al. (1986) regard Noturus insignis as native to the 
Monongahela drainage. 

Noturus insignis has entered the Mississippi and 
Great Lakes drainages by relatively recent stream 
changes or human introduction. Changes in 
drainage pattern may account for dispersal into the 
Holston drainage but apparently do not account 
for its appearance in other drainages. 

Madtoms are often used as bait by anglers. Since 
they are hardy and able to survive injury, disposal 
of unwanted bait specimens could have started new 
populations in areas with favourable habitat. The 
Margined Madtom has undoubtedly been 
introduced into Clark Lake, Gogebic County, 
Michigan (UMMZ 186551) and presumably into 
the Merrimack River of New Hampshire (Taylor 
1969). Schmidt (1986) suggests the Merrimack 
drainage population of Noturus insignis could be 
native but since there are no recorded populations 
between the Merrimack and Hudson he suggests 
the argument is less convincing than for Noturus 
gyrinus, which has a similar distribution. 

One of the ecological factors which seems to 
contribute to the present distribution of Noturus 
insignis is the availability of well-oxygenated water 
sufficiently warm for reproduction. Although 
Noturus species commonly avoid cold water, 
Noturus insignis may occur in salmonid streams 
indicating a higher tolerance for cold water. 

Canada: Since Noturus insignis was not found 
in Canadian waters prior to 1971, it was excluded 
from the first edition of Freshwater Fishes of 
Canada by Scott and Grossman (1973) but is 
mentioned briefly in the notes of the 1979 reprint. 

In August 1971, four specimens were collected in 
a stream draining Lac a la Loutre to Lac Lapeche, 
in Gatineau Park, Quebec, 45 km northwest of 



Ottawa. An additional 25 specimens were collected 
at the same location in November 1971 (Figure 3). 
This location was more than 20 1 km ( 1 25 air miles) 
north of its previously known limit in New York 
State (Rubec and Goad 1974). These collections 
are catalogued at the Canadian Museum of Nature 
(NMG), at Ottawa [NMC 73-0142, NMC 74-0004]. 
In May 1976, a single specimen of Noturus 
insignis was identified in an Ontario Ministry of 
Natural Resources (OMNR) collection from the 
Fall River, Lanark County, Ontario, 85 km 
southwest of Ottawa (Figure 3). Since this was the 
first reported occurrence of this species in Ontario, 











FuiURE 2. North American distribution ol the Mar- 
gined Madtom, Noturus insignis. Squares 
represent the disjunct Canadian populations 
{adapted from Lee et al. 1980). 



1990 



GOODCHILD: STATUS OF THE MARGINED MADTOM 



31 




Figure 3. Distribution of the Margined Madtom, 
Noturus insignis, in Ontario and Quebec. 



OMNR crews returned to the Fall River to 
determine the extent of a Margined Madtom 
population (G. A. Goodchild, Fisheries Branch, 
Ontario Ministry of Natural Resources, Queen's 
Park, Toronto; personal communication). On 
June 17, an additional 14 specimens were collected 
in the Fall River by electrofishing (one specimen 
was found dead on the banks). Thirteen of these 
fish are catalogued in the Royal Ontario Museum 
(ROM) collection [ROM32581]. 

Since there are well-documented introductions 
of Noturus insignis into streams in the United 
States, it is possible that its occurrence in Canada 
may also be explained by the release of baitfish by 
anglers. The two areas in Canada where Margined 
Madtoms have been found (Figure 4) are both 
popular areas for sport fishing and both are in or 
adjacent to large parks, Gatineau Park, in Quebec, 
and Silver Lake Provincial Park, in Ontario. 

Bailey and Smith (1981) felt that Noturus 
insignis entered the Great Lakes Basin by 
introductions and not by post-glacial dispersal or 
recent immigration through canals and water 
course alterations. Although Taylor (1969) 
suggested that Noturus insignis entered the Great 
Lakes drainage by relatively recent stream changes 
or human introductions, he hypothesized that the 
relatively northern Noturus flavus utilized post- 



glacial stream changes to enter the Great Lakes at a 
number of points. 

McAllister and Coad (1974) suggested that 
presence of Margined Madtom in Canada may 
represent either a relict population originating 
from the Atlantic Coastal plain, indicative of a 
formerly wider range, or an introduction. 
McAllister (1987) pointed out that Noturus 
insignis was now recorded in western New York 
state, only 130 km from the Canadian record in the 
Gatineau basin and suggested it is plausible that 
the Canadian occurrence is natural. Grossman and 
VanMeter (1979) considered that early records 
from small tributary streams in the Oswego River, 
New York and records from the tributary of the 
Ottawa River, could indicate the presence of 
isolated populations in the extreme eastern part of 
the Lake Ontario watershed. Margined Madtom 
have been reported from Oswego Country, New 
York as recently as 1985 (McAllister 1987). 

During the final retreat of the Wisconsin ice 
sheets from the Gatineau area, the Champlain Sea 
overlaid the area of the present Gatineau, Rideau, 
Ottawa, and St. Lawrence rivers. Lake Champlain 
and the Hudson River. Although the initial stages 
of the Champlain Sea were marine, evidence 
indicates a fresh-water drainage was initiated 
about 10 000 years ago (Romanelli 1975; 
McAllister and Coad 1974). Lake Frontenac was 
of relatively short duration in the Ottawa area but 
this postglacial lake was confluent with glacial 
Lake Iroquois (Lake Ontario) and glacial Lake 
Vermont (present Lake Champlain). It provided a 
corridor by which dispersal could have taken place 
in the Ottawa Region (Rubec 1975). Also, rapid 
warming conditions at that time led to a warmer 
cUmate than exists today, allowing warm water 
species into the region by way of the Ottawa River 
(Rubec 1975). The final connection between the 
Ottawa River and the Great Lakes disappeared 
4000 years ago and from that time until the 
construction of the Rideau Canal in the early 
1800s, dispersal was restricted to ascending the 
Ottawa River (McAllister and Coad 1974). 

Extremely limited population numbers may cast 
some doubt on the theory that Noturus insignis 
became established in the Ottawa River drainage 
during post-glacial events. These numbers may, 
however, be a reflection of inadequate suitable 
habitat. Similar disjunct records are known from 
New York State and New Hampshire. Addition- 
ally, the paucity of Ottawa records may reflect the 
lack of suitable collecting techniques. 

The possibility that the Margined Madtom is a 
canal immigrant cannot be completely ignored. In 
New York State, the construction of many canals 
has considerably altered the distribution of fish 
species within the state (Smith 1985). Disjunct 



32 



The Canadian Field-Naturalist 



Vol. 104 




Figure 4. Distribution of Noturus insignis (eastern Ontario and western Quebec) 



distribution points in the St. Lawrence River 
drainage of New York State are indicated on the 
distribution maps for Margined Madtom in Smith 
(1985) and Lee et al. (1980), but not explained 
(Figure 2). Recent dispersal of Margined Madtom 
to other new locations, such as their entry into the 
East River in West Virginia, is well documented 
(Stauffer et al. 1975). Yet, the waterfall at 
Wakefield on the lower Riviere Lapeche appears to 
be an effective barrier to the dispersal of Ottawa 
River fish fauna into Gatineau Park (Rubec 1975), 
which argues against the Rideau Canal dispersal 
hypothesis. There is little direct evidence for any 
uniqueness of the populations in Canada. The 
differing views of origin will only be settled if 
additional populations are discovered, particularly 
in areas remote from sport fishing (Coad 1986), or 
if evidence is found of genetic differentiation in 
Canadian specimens. Given the restrictions on the 
use of bait fish in Ontario and Quebec and the 
elusive character of the species, however, it is 
doubtful that the species was introduced here and 
one should tentatively assume that it is an 
indigenous species. 



Protection 

No specific protection exists in Canada other 
than that afforded by the habitat sections of the 
Fisheries Act. Noturus insignis is not a legal bait 
species in Ontario and the use of bait is illegal in 
Quebec. 

Population Sizes and Trends 

Only 49 individuals have been captured in 
Canada. The majority were collected between 1971 
and 1976. 

The original locations where Margined 
Madtom were taken in Quebec were re-sampled 
in 1982, 1983 and 1984 in an attempt to assess the 
status of this species. No individuals were 
captured or observed at these sites, but a single 
fish was caught upstream in June 1982 [NMC 82- 
0321], and a second was caught and released from 
the same location in September 1982. Two 
specimens were caught in Riviere Lapeche near 
Saint-Louis-de-Masham on I September 1982 
[NMC 82-0572] and another on 19 July 1983 
[NMC 83-0S35]. Riviere Lapeche drains Lac 
Lapeche and lies in the same drainage as previous 
Quebec collections. Extensive sampling in the 



1990 



GOODCHILD: STATUS OF THE MARGINED MADTOM 



33 



area (Gatineau Park) did not reveal any other 
localities for Noturus insignis (Coad 1986). 

In Ontario, Coad (1986) reports that attempts to 
capture the Margined Madtom in the Fall River, 
Lanark County, in July 1982 were unsuccessful 
despite exhaustive sampling of riffle areas. This 
area was originally sampled by OMNR as part of 
the Stream Inventory Program (qualitative 
sampling program used to provide base-line 
assessment of streams in the Province) (G.A. 
Goodchild, personal communication). 

Madtoms are notoriously difficult to capture. 
According to Taylor (1969), species of Noturus are 
mainly active at night and hide during the day. In 
fact, one species in the United States was collected 
and described but could not subsequently be 
collected from the type locality for many years, 
despite attempts by a plethora of renowned 
ichthyologists (B. Coad, Curator, National 
Museum of Natural Sciences, Ottawa, Ontario; 
personal communication). 

Some species of Noturus are quite irregularly 
distributed and infrequently captured except by 
intensive survey work. Since Madtoms tend to be 
secretive, they are most readily collected by 
chemicals, electrofishing or seining after dusk 
(Taylor 1969). Madtoms may often be present in 
quite low numbers. Populations fluctuate wildly 
and are dependent on spawning success from year 
to year (Trautman 1981; Bauer et al. 1983), 
therefore it is difficult to assess their numbers and 
status. Coad (1986) suggests the initial collecting 
expeditions may have even contributed to 
depletion of Margined Madtom populations in 
Canada. Perhaps they still remain in the two main 
areas in Canada, but are uncommon and variable 
in year class success (B. Coad, personal 
communication). 

There is insufficient evidence to evaluate 
population structure of Noturus insignis. Because 
of its restricted distribution, low population 
numbers and limited habitat, the continued 
survival of this species in Canadian waters may be 
doubtful. It is now either absent from the two 
known localities or present in numbers reduced 
below those necessary to ensure capture (Coad 
1986). 

Habitat 

The Margined Madtom inhabits clearwater 
streams of moderate current. It is usually found in 
riffles where it lives among rocks, boulders or 
coarse gravel (Hubbs and Lagler 1967; Taylor 
1969; Lee et al. 1980; Smith 1985). Taylor (1969) 
also states that Noturus insignis are recorded from 
high-gradient streams above the Fall Line. 

In Canada, McAllister and Coad (1974) report 
that the Margined Madtom is a fluviatile species 



found chiefly in high gradient streams. It lives in 
clean or normally clear water in moderate to swift 
current, about riffles, rubble, boulders, or coarse 
gravel. The first specimens reported from Canada 
were said to inhabit a situation typical for the 
species in the United States (Rubec and Coad 
1974). 

The extremely specific habitat requirements of 
the Margined Madtom are well documented. It 
does not appear to be able to exist in even slightly 
divergent circumstances, despite its evident 
hardiness. 

Exhaustive re-sampling at the first sites of 
capture in Canada failed to produce additional 
captures although specimens have been caught 
upstream as well as in a nearby stream. There has 
been an increase in Beaver {Castor canadensis) 
activity along the stream creating a series of dams 
that have submerged the original riffle areas (Coad 
1986). Coad (1986) reports that the water flow was 
slower and silt was found covering the pebbles and 
cobbles. He suggests that loss or fluctuations in 
suitable conditions may severely affect the survival 
of a population limited in numbers. The area may 
no longer be suitable for the Margined Madtom. 

General Biology 

Reproductive Capability: There have been no 
reported studies on the life history of this species in 
Canada. No observations of spawning activities 
were recorded in Canada although collections were 
made during the period of time when they have 
been observed spawning in north-eastern United 
States. 

Little is known about the spawning of Noturus 
insignis in the United States. The species is 
nocturnal and probably spawns in relative 
darkness, under cover (Taylor 1969). Eggs are 
deposited under rocks near small rapids in a quiet 
extent of water. They are likely soUtary spawners. 
Males were observed swimming passively near the 
nests by Fowler (1917). Bowman (1936) observed 
spawning in late June, while Clugston and Cooper 
(1960) found that all females collected were spent 
by 30 July. Almost all were spent by the middle of 
July, indicating that spawning occurred in late 
June to early July in Pennsylvania. 

Females appear to be mature in their third 
summer or at age II, some males may attain sexual 
maturity in their second summer, a year before the 
females (Clugston and Cooper 1960). 

All madtoms lay comparatively few eggs, many 
species less than 100 at a time (Taylor 1969). 
Clugston and Cooper (1960) gave an egg count of 
107 from a 122 mm female and Fowler (1917) 
estimated 200 eggs per nest. 

Species of Noturus are said to be characterized 
during breeding season by swollen areas on the 



34 



The Canadian Field-Naturalist 



Vol. 104 



head and upper surfaces and by a drab colour 
(Taylor 1969). Fowler (1917) describes the 
conspicuous guardian male of Noturus (Schil- 
beodes) insignis as having pale barbels and pale fin 
edges. 

However, Clugstonand Cooper (1960) found no 
reUable external sex characters and determined sex 
by dissection. During the spawning period, 
collections had a significant deviation from the 
50:50 sex ratio found during other periods but 
these deviations were not Ukely due to male 
mortality. These authors felt that since males 
guard the nests and young (Fowler 1917; Bowman 
1932), they were less readily captured. 

Population structure was studied by Clugston 
and Cooper (1960) in Pennsylvania. They found 
that most of the fish collected were in age-groups I, 
II and III with a few in groups and IV. The oldest 
fish was four years old. They were up to 158 mm in 
total length (TL) and a specimen 136 mm TL 
weighed 26.5 g. During the first two years, annual 
increments of growth averaged 45 mm, while in 
third and fourth years, increments decreased to 
about 20 mm. In both sexes, there is a high growth 
rate during the summer months. After sexual 
maturity is reached, most of the growth occurs in the 
warm months after spawning, indicating the 
majority of food taken by mature fish up to the time 
of spawning was used for egg/ sperm production and 
after spawning, it was used for growth. 

Behaviour I Adaptability: Since the Margined 
Madtom is a secretive, nocturnal species, it is not 
readily observed and therefore little is known 
regarding the effects of human interference. It has 
been used as a bait fish, particularly in 
Pennsylvania (Rubec and Coad 1974), and is 
evidently hardy and able to withstand transport as 
indicated by documented range extensions. 

Reported foods include cladocerans, ostracods, 
gammarids, midges and debris (Smith 1985). 
Insects and unidentified fish were found in three 
individuals by Flemer and Woolcott (1966). 

Limiting Factors 

Noturus insignis is restricted to moderate to 
rapidly flowing streams with an abundance of 
cover. Although wet specimens will survive for 
several hours in air, oxygen deficient water appears 
to be a critical factor in controlling distribution. In 
late summer, madtoms become almost entirely 
restricted to riffles when oxygen is depleted in 
pools (Taylor 1969). 

Coad ( 1 986) felt that stream alterations resulting 
from Beaver dams were partly responsible for the 
depletion of the population in a stream in Gatineau 
Park. Any barrier, obstruction or activity that 
reduced riffle areas would undoubtedly have a 
severe effect on populations of the Margined 
Madtom. 



As mentioned by Coad (1986) and documented 
by Trautman (1981) and Bauer et al. (1983), 
populations of species of Noturus may fluctuate 
depending on spawning success. Collection of 
specimens may also contribute to depletion of 
populations already low in numbers. 

Special SigniHcance of the Species 

The presence of Noturus insignis in Canada is an 
interesting anomaly. It is one of the freshwater 
madtoms which can inflict a painful, but not 
dangerous, wound from the pectoral spines and 
associated poison gland (Hubbs and Raney 1944; 
Scott and Crossman 1973). The species is of no 
economic importance and its continued existence 
in Canada is debatable. Its presence here is of 
considerable interest to an understanding of the 
glacial history of the area and the species could 
provide information on the evolutionary processes 
in madtoms. 

Evaluation 

The Margined Madtom populations in Ontario 
and Quebec are found over 130 km north of the 
United States records. There are two possible 
explanations for the occurrence of this species in 
Canada: (1) Margined Madtoms may have been 
introduced to Canadian waters by the accidental 
deposition of baitfish and have managed to survive 
in restricted areas in very low numbers; or (2) 
Margined Madtoms have been continuously 
present since post-glacial dispersal, surviving in 
isolated populations in the eastern part of the 
Great Lakes watershed (Crossman and VanMeter 
1979). 

Given the restrictions on the use of bait fish in 
Ontario and Quebec and the glacial history of the 
area, the latter hypothesis is more plausible. Since 
it survives in low numbers and in restricted areas; is 
at the fringe of its natural range and thus is in 
jeopardy in Canada, it should be classified as 
threatened. 

Acknowledgments 

The assistance of D. E. McAllister and B. W. 
Coad of the National Museum of Natural Sciences 
(now Canadian Museum of Nature); E. Holm, 
R. W. Winterbottom and E. J. Crossman of the 
Royal Ontario Museum; G. E. Gale and G. A. 
Goodchild of the Ontario Ministry of Natural 
Resources in providing access to records and 
reports was invaluable. The author also wishes to 
thank R. R. Campbell, Fisheries and Oceans for 
his helpful comments on the manuscript and 
support in the preparation of this report. Financial 
assistance in the preparation of this report was 
provided by World Wildlife Fund (Canada). 



1990 



GOODCHILD: STATUS OF THE MARGINED MADTOM 



35 



D. E. McAllister gave permission to use the 
drawing of Noturus insignis from the book Fishes 
of Canada 's National Capital Region. 

Literature Cited 

Bailey, R. M., and G. R. Smith. 1981. Origin and 
geography of the fish fauna of the Laurentian Great 
Lakes basin. Canadian Journal of Fisheries and Aquatic 
Science 38: 1539-1561. 

Bauer, B. H., G. R. Dinkins, and D. A. Etnier. 1983. 
Discovery of Noturus bailey i and N . flavipinnis in Citico 
Creek, Little Tennessee River System. Copeia 1983(2): 
558-560. 

Bowman, H. B. 1936. Further notes on the Margined 
Madtom Rabida insignis and notes on a kindred species 
Noturus flavus Rafinesque. Ph.D Thesis, Cornell 
University. 

Clugston, J. P., and E. L. Cooper. 1960. Growth of the 
common eastern madtom, Noturus insignis, in central 
Pennsylvania. Copeia 1960 (1): 9-16. 

Coad, B. W. 1986. The Margined Madtom [Noturus 
insignis) in Canada. Trail and Landscape 20(3): 
102-108. 

Grossman, E. J., and H. D. VanMeter. 1979. 

Annotated list of the fishes of the Lake Ontario watershed. 
Great Lakes Fishery Commission Technical Report 36: 
1-25. 

Flemer, D. A., and W. S. Woolcott. 1966. Food habits 
and distribution of the fishes of Tuckahoe Creek, 
Virginia with special emphasis on the bluegill, Lepomis 
m. macrochirus Rafmesque. Chesapeake Science 7(2): 
75-89. 

Fowler, H. W. 1917. Some notes on the breeding habits 
of local catfishes. Copeia 42: 32-36. 

Hocutt, C. H., R. E. Jenkins, and J. R. Stauffer, 
Jr. 1986. Zoogeography of the fishes of the Central 
Appalachians and Central Atlantic Coastal Plain. 
Pages 161-211 in The zoogeography of North 
American freshwater fishes. Edited by C. H. Hocutt 
and E. O. Wiley. John Wiley and Sons, New York, 
New York. 866 pages. 

Hubbs, C. L., and E. C. Raney. 1944. Systematic notes 
on North American siluroid fishes of the genus 
Schilbeodes. Occasional Paper, Museum of Zoology, 
University of Michigan 487. 36 pages. 

Johnson, J. E. 1987. Protected fishes of the United 
States and Canada. American Fisheries Society, 
Bethesda, Maryland. 42 pages. 

Lee, D. S., G. R. Gilbert, G. H. Hocutt, R. E. Jenkins, 
D. E. McAllister, and J. R. Stauffer, Jr. 1980. Atlas 
of North American freshwater fishes. North Carolina 
State Museum of Natural History Biological Survey 
Publication 1980-12. 867 pages. 



McAllister, D. E. 1987. Review of: The inland fishes of 

New York State by C. L. Smith. 1986. Canadian 

Field-Naturalist 101: 646-647. 
McAllister, D. E., and B. W. Goad. 1974. Fishes of 

Canada's National Capital Region. National Museum 

of Natural Sciences Miscellaneous Special Publication 

24. 200 pages. 
Miller, R. R. 1972. Threatened freshwater fishes of the 

United States. Transactions of the American Fisheries 

Society 101(2): 239-252. 
Romanelli, R. 1975. The Champlain Sea episode in the 

Gatineau River Valley and Ottawa area. Canadian 

Field-Naturalist 89(4): 356-360. 
Rubec, P. J. 1975. Fish distribution in Gatineau Park, 

Quebec, in relation to postglacial dispersal, man's 

influence, and eutrophication. Canadian 

Field-Naturalist 89(4): 389-399. 
Rubec, P. J., and B. W. Goad. 1974. Fist record of the 

Margined Madtom, Noturus insignis, from Canada. 

Journal of the Fisheries Research Board of Canada 

31(8): 1430-1431. 
Schmidt, R. E. 1986. Zoogeography of the Northern 

Appalachians. Pages 137-159 in The zoogeography of 

North American freshwater fishes. Edited by C. H. 

Hocutt and E O. Wiley. John Wiley and Sons, New 

York, New York. 866 pages. 
Scott, W. B., and E.J. Grossman. 1973. Author's 

comments in Freshwater Fishes of Canada 1979 

Revised Printing. Fisheries Research Board of Canada 

Bulletin 184. 966 pages. 
Smith, G.L. 1985. Theinlandfishesof New York State. 

New York State Department of Environmental 

Conservation. Albany, New York. 522 pages. 
Stauffer, J. R., Jr., G. H. Hocutt, M. T. Masnik, and 

J. E. Reed, Jr. 1975. The longitudinal distribution of 

the fishes of the East River, West Virginia-Virginia. 

Virginia Journal of Science 26(3): 121-125. 
Taylor, W. R. 1969. A revision of the Catfish genus, 

Noturus, Rafinesque, with an analysis of the higher 

groups in the Ictaluridae. U.S. National Museum 

Bulletin 282. 315 pages. 
Trautman, M. B. 1981. The fishes of Ohio with 

illustrated keys. Revised edition. Ohio State University 

Press, Columbus, Ohio. 782 pages. 

Accepted 10 October 1989 

Addendum 

Since this report was completed, two additional 
specimens of Noturus insignis have been collected 
from Lake Joseph, Muskoka District, Ontario, by 
OMNR in 1988 and 1989 (ROM 55829; 58161). 



Status of the Brook Silverside, Labidesthes sicculus, in Canada* 
Cheryl D. Goodchild 

2168 Harcourt Crescent, Mississauga, Ontario L4Y IWl 

Goodchild, Cheryl D. 1990. Status of the Brook Silverside, Labidesthes sicculus, in Canada. Canadian Field- 
Naturalist 104(1): 36-44. 

The Brook Silverside, Labidesthes sicculus, is not in any COSEWIC category. Confined to a small region in south- 
central Canada, populations of Brook Silverside are contiguous with those in the United States, but are at the northern 
hmit of the species' range. Although there have been no population studies, collection records do not indicate any 
evidence of a decline. Populations appear to be relatively stable and could be expanding slightly. Somewhat tolerant of 
alterations in habitat, it is extremely sensitive to turbidity and interspecific competition. This coupled with its one-year 
life cycle has the potential of completely decimating populations in a very short time span. Therefore, regular 
monitoring of this species should be encouraged. 

Le crayon d'argent, Labidesthes sicculus, ne figure dans aucune categoric du CSEMDC. Confinee dans une aire 
restrainte du centre sud du Canada, cette espece dont les populations sont voisines de celles des Etats-Unis, occupe la 
limite septentrionale de son territoire. Bien qu'aucune etude demographique n'ait encore ete effectuee, les donnees sur 
les prises en revelent aucun signe de regression. Les populations semblent assez stables et pourraient meme etre 
legerement en expansion. Malgre sa tolerance relative aux transformations de son habitat, le crayon d'argent est 
extremement vulnerable a la turbidite et a la competition interspecifique, ce qui, outre sa duree de vie d'un an, pourrait 
conduire a la disparition de populations entieres en un laps de temps tres court. En consequence, il faut encourager la 
surveillance reguliere de cette espece. 

Key Words: Brook Silverside, Labidesthes sicculus, crayon d'argent, Atherinidae, silversides, freshwater fishes, status, 
Canada. 



The Brook Silverside, Labidesthes sicculus 
(Cope 1865), is the only member of the family 
Atherinidae that inhabits Canadian freshwaters, 
although the marine Atlantic Silverside {Menidia) 
is sometimes found in brackish waters (Scott and 
Grossman 1973; Scott and Scott 1988). This is a 
large family containing mostly marine fishes, only 
two species occur in freshwater in North America; 
the Inland Silverside {Menidia beryllina) in the 
United States only and the Brook Silverside in 
both Canada and the United States (Cooper 1983). 

Labidesthes sicculus (Figure 1) a small, slender, 
somewhat pellucid fish is described as straw, pale 
green or olive coloured. The most conspicuous 
feature is the brilliant silvery lateral band. Since 
the body is rather transparent, the swim bladder 
and vertebral column can be seen through the 
muscle and peritoneum, particularly in small fish 
(38 to 51 mm) [Scott and Crossman 1973; 
McAllister and Coad 1974]. Adults average 76 mm 
(3 in) in total length (TL), but have been reported 
up to 1 10 mm (4.4 in) in Pfieiger (1975). 

The inconspicuous first dorsal fin may have 
from three to six (four in Canadian specimens) 
weak spines. Both dorsal fins are positioned over 
the extremely large sickle shaped anal fin which 
has one spine and between 23 to 27 rays. The top of 



the head is flattened and the snout is long and 
pointed. The upper jaw is quite protrusible and is 
projected into a short beak while the mouth is 
relatively large. Brook Silversides do not develop 
breeding tubercles nor do they have special 
breeding colours (Scott and Crossman 1973; 
Cooper 1983; Smith 1985). 

Distribution 

North America: Widely distributed throughout 
the freshwaters of central North America, the 
Brook Silverside is found in the Great Lakes, 
Mississippi, and Gulf Coast drainages (Cooper 
1983). On the Atlantic coast, it ranges north from 
Florida to South Carolina (Smith 1985). At one 
time two forms were recognized, one thought to 
occur in the lower Mississippi Valley, Gulf Coast 
and Atlantic Coast part of the range, and the other 
from the northern Mississippi Valley and Great 
Lakes part of the range (Hubbs and Lagler 1967; 
Lee 1980). However, Bailey et al. (1954) 
recommended they not be recognized as subspe- 
cies. Also, a recent study provides the first 
published karyotypes for North American 
atherinids. The subspecies of Labidesthes sicculus 
were examined for chromosomal differences but 
none were noted (Korth and Fitzsimmons 1987). 



♦Received and accepted by COSEWIC 1 1 April 1989 no designation required. 



36 



1990 



GOODCHILD: STATUS OF THE BROOK SiLVERSIDE 



37 



g7 ' ^?^^.^'-v " ■ "; - - ■- '"-. ■ '" '■ -■ ■ . ^ ^w - ny. ' v .- ■' ■ ^fe^^iiff iP j i j ^;^,^ 




Figure 1. Drawing of the Brook Silverside, Labidesthes sicculus (drawing by A. Odum; from Scott and 
Grossman (1973) by permission). 



In the United States, the Brook Silverside is 
found in the Allegheny River system of New York 
state, west to the lower peninsula of Michigan 
through Wisconsin and Minnesota, south to the 
Gulf states of Texas and Oklahoma, along the Gulf 
coast to peninsular Florida, and north on the 
Atlantic coast as far as South Carolina (Figure 2) 
[Scott and Grossman 1973; Lee 1980; Smith 1985]. 

It was first reported from the Lake Superior 
drainage in 1975 when it was collected in the Dead 
River near Marquette, Michigan, both at the 
mouth and 100 m upstream (Berg et al. 1975). 
However, Cahn (1927) had placed it in the 
headwaters of the Upper Fox River and Shawano 
Lake, Lake Michigan watershed and also in Chain- 
0-Lakes, northwestern Washburn County, 
Mississippi River watershed, all in Wisconsin. 

Lack of prior records in the Lake Superior 
drainage suggest that it is indigenous, but very 
rare, or that it has recently appeared, perhaps by 
introduction or by migration through the locks at 
Sault Ste. Marie (Berg et al. 1975). Several new 
species have appeared in the Lake Superior 
drainage in the last 10 years due to ballast water 
introductions (G. A. Goodchild, Fisheries Branch, 
Ontario Ministry of Natural Resources, Toronto, 
Ontario; personal communication). 

There are few reported range extensions for this 
species and generally it seems to be declining in the 
United States. Before 1900, in Ohio, Brook 
Silverside were common in large streams such as 
the Maumee, Scioto, and possibly the Ohio River, 
but today are almost absent in these now turbid 
waters (Trautman 1981). They were formerly 
present in the Youghiogheny River, in southwest- 
ern Pennsylvania, but are now limited to northwest 
parts of the state (Cooper 1983). In New York 
State, it was believed to be a canal immigrant to the 
Mowhawk River but records from the 1920s to 
1930s have not been duplicated in recent years 
(Smith 1985). Although evidently disappearing in 
some states, the Brook Silverside is still common in 
the Lake Erie drainage, western Allegheny 
drainage and the Finger Lakes. It is particularly 
common in the old Erie Canal of New York State. 



Canada: The Brook Silverside is found in the 
drainages of the upper St. Lawrence River, the 
lower Ottawa River, in the drainage basins of lakes 
Ontario, Erie, St. Clair, and Georgian Bay (Scott 
and Grossman 1973). 

The distribution of Brook Silverside in Canada 
(Figure 3) has changed httle since first documented 
by Radforth (1944). It has probably expanded in 
the northern part of its range. The species is most 
likely indigenous and probably arrived in southern 
Ontario (Figure 4) and Quebec via a number of 
postglacial dispersal routes. The Mohawk- 
Hudson, Fort Wayne and possibly the Champlain 
and Chicago outlets were suggested by Radforth 
(1944) to have been routes used to enter Ontario. 
Brook Silverside probably used the Chicago and 
Grand Valley outlet (Mandrak 1990) to disperse 
into lakes Huron and Erie. Dispersal into Lake 
Ontario may have occurred prior to the formation 
of Niagara Falls (approximately 12 500 B.P.), and 
dispersal into Georgian Bay probably occurred 
during the late glacial Lake Algonquin stage, 
providing access across the Bruce Peninsula (N. E. 
Mandrak, personal communication). They 
presumably survived glaciation in the Mississippi 
Valley refugium, showing a clear affinity for the 
Mississippi Valley fauna (Schmidt 1986). 

McAllister and Goad (1974) reported that the 
northern limit of the range of Labidesthes sicculus 
was reached in the National Capital Region 
(Rideau River). Subsequent surveys have collected 
specimens from the Ottawa River as far as Kettle 
Island Bay, south of Gatineau Park [Canadian 
Museum of Nature (NMC 79-1224, NMC 85- 
0608]. The most northerly collection of the Brook 
Silverside appears to be from Wilson Lake near 
Pointe au Baril Station, northwest of Parry Sound 
[Royal Ontario Museum (ROM) 30654]. Wilson 
Lake drains into Georgian Bay via Six Mile Lake 
and the Naiscoot River. This is an extension of the 
previously known range. 

In Georgian Bay it has been collected from Big 
Sound, Parry Sound, by the Ontario Ministry of 
Natural Resources (OMNR) in 1979. Bensley 
(1915) had found it to be common around the 



38 



The Canadian Field-Naturalist 



Vol. 104 




—49° 



-30° 



Figure 2. North American distribution of the Brook Silverside, Labidesthes sicculus 
(adapted from Lee 1980). 



shore in Georgian Bay and in tributaries such as 
the Go Home River. Recent collections from 
Woods Bay, Victoria Harbour, Matchedash Bay 
and Honey Harbour indicate the Brook Silverside 
is still quite common in southern Georgian Bay 
(OMNR and ROM records). 

Although presence in Lake Huron, proper, has 
not been mentioned in the literature it is not 
surprising that it has been found there due to its 
occurrence in both the St. Clair River and 
Georgian Bay. In 1986, specimens were taken 
inside the breakwall in Sarnia Bay of Lake Huron 
by OMNR collectors. It has also been collected in 
the Bayfield River, south of Goderich, Ontario, a 
tributary to Lake Huron [ROM 43028]. It is 
frequently found in many tributaries of Lake 
Huron (Ausable, Maitland, and Saugeen rivers). 
Lake Erie (Grand and North rivers), and Lake St. 
Clair (Sydenham and Thames rivers) [NMNS; 
OMNR; ROM]. 

In his studies of the fishes of eastern Ontario, 
Toner (1943) found it was common in both Lake 
Ontario and the St. Lawrence River but recorded it 
from only one inland lake (Grippen Lake, of 
central Leeds County). Elsewhere in Leeds 



County, he considered it was either rare or absent. 
In contrast, Scott (1967) considered Brook 
Silverside to be very abundant in some inland lakes 
in Leeds County, particularly in summer. It has 
since been collected in Grippen Lake in the early 
1970s as well as several other lakes in the area: Mud 
Lake, Mosquito Lake, Opinicon Lake, and 
Whitefish Lake (OMNR and ROM records). 

Protection 

General protection is afforded by habitat 
sections of the Fisheries Act, but there is no specific 
protection for the species in Canada. Since it is not 
classed as a baitfish in Ontario, it cannot be sold as 
such so this would provide some protection. 

In the USA, it was listed as rare in Maryland and 
Pennsylvania (Miller 1972). It is not included in the 
American Fisheries Society's Protected Fish of the 
United States and Canada (Johnson 1987). 

Population Sizes and Trends 

Labidesthes sicculus is a common forage fish in 
lakes and streams of eastern North America. Cahn 
(1927) described it as being one of the most 
abundant and typical species in Waukesha 



1990 



GOODCHILD: STATUS OF THE BROOK SILVERSIDE 



39 




Figure 3. Distribution of Brook Silverside, Labidesthes sicculus, in Canada. 



County, Wisconsin and considered it to be very 
abundant throughout much of its range in the 
United States. Brook Silverside were usually the 
most abundant species in seine hauls of Crooked 
Lake, Indiana (Nelson 1968). 

There are few reported range extensions for this 
species and populations generally seem to be 
declining. In Illinois, it is somewhat decimated due 
to excessive siltation (Smith 1979). Cooper (1983) 
describes it as being more common in lakes than in 
streams in Pennsylvania and although formerly 
present in the Youghiogheny River, it is now 
extremely limited to the northwestern parts of the 
state. 

Trautman (1981) discusses extensively the 
decline in Brook Silverside in Ohio. Before 1900, it 
was usually found to be abundant in most areas 
investigated. In surveys undertaken in the 1930s to 
1950s, it was far less abundant, taken in less than a 
tenth of the collections made. Since the 1940s, 
Brook Silverside have been found primarily in 
small clear upland brooks or in recently 
constructed reservoirs. 

In contrast to the general trend of decreasing 
populations, Labidesthes sicculus seems to readily 



invade reservoirs and impoundments, quickly 
becoming abundant, particularly in those with 
clear water and sandy/gravel bottoms (Cross 1967; 
Smith 1979). In Missouri and Iowa, it is the most 
abundant small fish in large reservoirs (Harlan and 
Speaker 1956; Pflieger 1975). 

It appears that populations of Brook Silverside 
in impoundments may be extremely unstable and 
subject to collapse in a very short time. McComas 
and Drenner (1982) published the results of a study 
to determine the cause of sudden decrease in 
Labidesthes sicculus population in impound- 
ments. They found that populations are com- 
pletely replaced when in competition with Menidia 
beryllina, the Inland Silverside. 

In Canada, few observations indicate that the 
Brook Silverside attains an abundance compara- 
ble to that in lakes of the northern United States 
(Scott and Crossman 1973). For example, in 
Indiana at least 1 1 500 specimens were used in a 
study by Nelson (1968). In Grippen Lake, Leeds 
County, eastern Ontario, Toner (1943) noted that 
immense numbers could be seined at night on 
sandy beaches. He also observed large numbers in 
Lake Ontario, from a dock in Kingston. Bensley 



40 



The Canadian Field-Naturalist 



Vol. 104 




Figure 4. Distribution of Brook Silverside, Labidesthes 
sicculus, in Ontario (adapted from Radforth 
1944). 

(1915) reported that he observed enormous numbers 
of young Brook Silverside in schools offshore but 
commonly only collected a few specimens in seine 
hauls. 

In a study comparing growth of nine co-occurring 
fish species in Lake Opinicon, Ontario, at least as 
many specimens of Brook Silverside were captured 
and analyzed as other species but there is no indi- 
cation whether that is indicative of the population 
numbers present in the lake (Keast and Eadie 1984). 

Numbers of specimens, collected in various 
locations throughout Ontario since the early 1970s 
range from one to over 100, but are not indicative of 
exceptionally large populations. Since no population 
studies have been done on this species in Canada, 
little can be deduced regarding present population 
trends. 

Habitat 

The Brook Silverside is a surface-dwelling fish 
with a strong schooling tendency (Smith 1979). 
Preference for near surface water is characteristic of 
the species. The most outstanding feature of 
environmental preference is the remarkable 
difference in habitat selection displayed by juveniles 
and adults. Young Brook Silverside are one of the 
few freshwater fishes to adopt a pelagic surface 
habitat in large lakes soon after hatching (Cooper 
1983). Hubbs (1921) first reported this phenomenon 
and he found the tendency more pronounced in lakes 
than in streams. Young Brook Silverside prefer to 
occupy the top few centimeters over deep water. In 
contrast, the adults show an exclusive selection of 
shallow water over shoals. Young fish move to a 
permanently shallow water habitat after attaining 
approximately two thirds of adult size (Cahn 1927). 
This occurs during August and September in 
Michigan and during winter the entire population 
can be found close to the shore beneath the ice 
(Hubbs 1921 after Evermann and Clark 1920). 



Brook Silverside are reported from widely 
variable habitats summarized by Lee (1980: 557) as 
follows: "abundant near surface of clear, vegetated 
and unvegetated warm waters of streams, lakes and 
reservoirs". In New York State it is reported to be 
most abundant in weedy areas of streams and lakes 
(Smith 1985). Although generally preferring areas 
with no noticeable current, it is occasionally taken 
from flowing water (Harlan and Speaker 1956). 
Tolerance of fairly rapid flow is also aptly 
demonstrated by collections from streams having 
considerable current, such as near the falls in the Go 
Home River (Bensley 1915). 

Although reasonably adaptable, the species is 
decidely intolerant of turbidity and is invariably 
absent from turbid waters. Trautman (1981) 
suggests it has moved into the small clear upland 
brooks or into cleaner backwaters of recently 
constructed reservoirs to avoid the turbid conditions 
now found in the large streams in Ohio, which it 
occupied prior to 1940. Brook Silverside also stop 
feeding when water becomes temporarily turbid 
(Trautman 1981; Smith 1985). However, specimens 
are often taken from turbid water in the Barge Canal 
System, New York State. 

Discrepancies to the reported substrate prefer- 
ence by Brook Silverside may be explained by 
choice of selectively different spawning habitat. 
Shallow water with an absence of vegetation but 
having sandy, gravelly, or rocky bottom is said to be 
characteristic habitat of the species by many authors 
(Toner 1943; Cross 1967; Trautman 1981). Yet, 
preference for localities with aquatic vegetation is 
described by others (Bensley 1915; Smith 1985). 
Possibly the species selects weedy areas only during 
spawning and hence those who reported a selection 
for aquatic vegetation had observed the species 
during the spawning period only. Spawning in 
spring among aquatic vegetation is reported by 
Scott (1967). Rooted aquatic vegetation was 
abundant in the section of the river where Berg et al. 
(1975) located Brook Silverside. Their collections 
were made in June and the specimens they kept were 
gravid females indicating that spawning was 
imminent. 

Although the distribution of Brook Silverside in 
Canada is limited, there is little evidence to suggest 
the species has declined appreciably since the early 
1900s when collections were first made. There have 
been no specific studies to determine the extent of 
Labidesthes sicculus populations in Canada and all 
the inherent difficulties associated with making 
estimates based on sporadic, unspecific sampling 
preclude determining critical habitat trends. 

General Biology 

Reproductive Capability: Labidesthes sicculus 
spawns at the age of one year. Although Fogle 



1990 



GOODCHILD: STATUS OF THE BROOK SILVERSIDE 



41 



(1959) noted a few males in Arkansas with two 
annuli, the oldest individuals collected in Crooked 
Lake, Indiana were estimated to be between 21 and 
23 months old and thus all spawning adults were 
oneyear of age (Nelson 1968). Hubbs (1921) found 
only one annulus evident on scales of breeding 
fishes taken in southeastern Michigan. Cahn 
(1927) examined 478 adults and never found more 
than one annulus. He concluded that Brook 
Silverside die before their second winter, probably 
at about 17 months of age. 

Spawning occurs in spring and early summer in 
and around aquatic vegetation. It can also occur 
over gravel in a moderate current and time of 
spawning may be temperature dependent. Due to 
incomplete data available in the literature, it is 
difficult to compare temperature/ time of spawn- 
ing at different locations. Cahn (1927) reported 
that pairing began when water temperature 
reached 18°C, actual spawning commenced at 
20° C and optimum spawning occurred at 23° C but 
he did not report exact dates. The breeding 
activities of Brook Silverside during May and June 
of a reportedly cool season in Michigan began after 
surface water temperature had risen above 20° C. 
Also, there was a temporary cessation of spawning 
when water temperature dropped below 20° C 
(Hubbs 1921). Indiana Brook Silverside spawn 
from the middle of June until early August but 
water temperatures are not reported (Nelson 
1968). Spawning in Missouri commences as early 
as 22 May but temperatures are also unavailable 
(Pflieger 1975). Labidesthes sicculus is identified as 
a protracted spawner in Lake Opinicon, Ontario 
with a duration of 41 days (11 June to 22 July) 
[Keast and Eadie 1984]. 

Little information is available on breeding 
frequency. Nelson (1968) observed fully ripe 
females containing mature orangish eggs up to 
1.2 mm but also noted were many immature 
whitish eggs up to 0.6 mm. This may indicate that 
females spawn more than once or merely that eggs 
may not all mature at once. Examination of 26 
females captured immediately after spawning 
found all but three with ovaries completely empty 
and these contained only half a dozen eggs each 
(Cahn 1927). Males on completion of a spawning 
act were seen pursuing other females. Possibly, 
males spawn repeatedly while females spawn only 
once. Males established ill-defined territories. 
Several males may chase a female moving into the 
area. Spawning occurs when one male and one 
female pair but no nest building or guarding occurs 
(Hubbs 1921; Cahn 1927; Nelson 1968) 

After extrusion, the egg settles slowly to the 
bottom. Each egg has at least one long adhesive 
filament which functions as an anchoring device 
for the non-adhesive egg. The number of filaments 



attached to Brook Silverside eggs may show a 
geographical gradation. Eggs from the Peace 
River, Florida, had two or three attachment 
filaments (Rasmussen 1980); eggs from Lake Fort 
Smith, Arkansas, had two filaments per egg (Fogle 
1959); while eggs from Portage Lake, Michigan 
(Hubbs 1921), Oconomowoc Lake, Wisconsin 
(Cahn 1927), and Crooked Lake, Indiana (Nelson 
1968), had only one filament. 

Scott and Crossman (1973) reported egg 
diameters of 0.8 to 1.2 mm but Rasmussen (1980) 
found egg diameters ranged from 1.1 to 1.4 mm. 
Mature eggs are orange in colour and contain 
numerous oil globules within the yolk. Eggs 
develop rapidly hatching in eight days in 25° C 
water and within eight to nine days in lake waters 
of 23°C(Cahn 1927). Rasmussen (1980) described 
the larvae and juveniles in considerable detail and 
notes geographical differences. 

No data are available on fecundity. In a study of 
Brook Silverside in Crooked Lake, Indiana, 
spawned eggs were considered to be numerous. 
However, year class strength may fluctuate 
considerably as evidenced by the much lower 
abundance observed during the second year of the 
two-year study (Nelson 1968). 

Growth of young Brook Silverside is rapid. 
Hubbs (1921) estimated an initial average growth 
of approximately a millimetre per day slowing to 
0.40 mm per day, attaining an estimated 70 to 80% 
of adult size by three to four months of age. Nelson 
(1968) noted that in an Indiana lake population, 
some fish reached the modal size of a 12-month-old 
spawning group in only three months. Rapid 
growth in the first months of life is critical to 
survival and attainment of sexual maturity in 
temperate freshwater fishes. Size-distribution 
histograms for nine species of fish in Lake 
Opinicon, Ontario, show that Brook Silverside 
were lighter per unit length than other species 
investigated, reflecting its long and narrow body 
shape (Keast and Eadie 1984). 

Species Movement: Long distance migration 
has not been observed for Brook Silverside. There 
is some evidence that the species may travel for 
some distance as it readily invades new impound- 
ments. Also, when river habitats become too 
turbid they readily move into clearer upstream 
areas. Large schools of Brook Silverside are 
frequently observed as well, particularly in lakes 
(Bensley 1915; Hubbs 1921). 

Labidesthes sicculus have a definite daily cycle 
of activity that seems to be regulated primarily by 
light intensity. Extremely active in the day, they 
usually become quiescent at night (Trautman 
1921). Being positively phototrophic, they are 
active on moonlit nights and night activity can be 
induced by shining a spot light on the water. 



42 



The Canadian Field-Naturalist 



Vol. 104 



Nighttime sampling may yield many more 
specimens than daytime collections (Cahn 1927). 
The use of artificial lights for fishing is not allowed 
in Ontario. 

Pelagic habitat selection by juveniles has already 
been discussed. There is also an interesting 
nocturnal-diurnal pattern of migration exhibited 
by young Brook Silverside. Prior to permanently 
adopting an inshore habitat in late August and 
September, young Brook Silverside migrate into 
littoral areas each night and return to the surface 
layer over deep water during the day. Cahn (1927) 
found that the inshore movement was coincident 
with a drop in temperature of the surface water 
over the depths as well as a change in the p H of the 
water at that stratum. By the end of the summer, all 
of the Brook Silverside become established inshore 
where they remain all winter. 

Behaviour I Adaptability: The Brook Silverside 
is a predacious fish that feeds on zooplankton and 
small insects (aquatic and terrestrial). It feeds in a 
snapping fashion with its beaklike toothed jaws 
and may frequently leap out of the water to catch 
flying insects (Scott and Grossman 1973; 
McAllister and Coad 1974). The diet of this highly 
specialized feeder is made up of Cladocera 
(frequently up to 80% by volume), small flying 
insects (up to 40%) and Chaoborus larvae (50%) 
[Keast and Webb 1966]. Occasionally other items 
may be utilized; Boesel (1938) reported finding a 
spider and small fish in the stomach contents of 
Brook Silverside examined. 

There is differential food selection between 
juveniles and adults. Boesel (1938) summarized the 
food items of two size ranges of Brook Silverside. 
Smaller individuals (average 35.9 mm) had a diet 
composed almost entirely of Entomostraca 
(mainly copepods), whereas larger individuals 
(average 61.2 mm) fed predominantly on insects, 
particularly adult midges. He found there was a 
gradation of feeding habits accompanying 
increasing size. The shift in diet from microcrusta- 
ceans to insects is concurrent with the migration of 
larger juveniles to a shallow water habitat (Pflieger 
1975). 

Parasites of Brook Silverside from Lake Erie 
have been recorded by Bangham and Hunter 
(1939) who noted that specimens from eastern 
Lake Erie were relatively unaffected although 
those from the west end of the lake harboured 
cestodes, trematodes, and nematodes. This was 
based on the examination of 30 species of which 10 
were infected with seven different species of 
parasites, in a latter study, Bangham (1972) 
resurveyed the parasites of Lake Eric and found 
four infected Brook Silverside out of the nine that 
were examined. All of these harboured species of 
trematodes. Hoffman (1967) lists five species of 



trematodes and one species of nematode for Brook 
Silverside. 

Labidesthes sicculus appears to be relatively 
tolerant of human disturbance readily invading 
newly created reservoirs and expanding into new 
areas when unfavourable habitat alterations occur 
in formerly occupied areas. It survives in areas 
where there has been considerable alteration of 
habitat through agriculture and construction as 
has occurred in southern Ontario. 

Limiting Factors 

Labidesthes sicculus is particularly susceptible 
to increased turbidity, a condition which has 
become more widespread in southern Ontario as a 
result of urbanization, construction and agricultu- 
ral activities (Scott and Grossman 1973). It 
disappears from waters that become turbid 
perhaps as a result of an inability to feed. Both 
Trautman (1981) and Smith (1985) report that 
activity and feeding cease when surface water 
becomes turbid. It has been suggested that 
cessation of feeding in turbid waters is triggered by 
the same mechanism (light intensity) that controls 
nocturnal inactivity. Brook Silverside are believed 
to feed actively mainly during daytime and rest 
during nighttime. However, Keast and Webb 
(1966) observed both nocturnal and diurnal 
feeding. 

Brook Silverside grow at an extremely rapid rate 
during their first few months of life. Therefore, any 
stress which might impede their ability to obtain 
sufficient food would have debilitating effects on 
the population. To determine the causes for rapid 
dwindling and extinction of populations of 
Labidesthes sicculus in a reservoir, a study of 
feeding mechanics was initiated by McGomas and 
Drenner (1982). They found that when in direct 
competition for zooplankton with another 
atherinid {Menidia beryllina). Brook Silverside 
experienced 75 to 86% mortality in laboratory 
studies. The two species have different feeding 
rates for copepods and in turbid water, feeding 
rates of Menidia were significantly higher, which is 
possibly another explanation for the strongly 
detrimental effect of turbid waters on Brook 
Silverside populations. 

Undoubtedly, Brook Silverside are extremely 
susceptible to catastrophic events due to their one 
year life cycle. If for example, an entire year class 
was decimated, then the complete population in 
that area could be destroyed. Brook Silverside may 
be restricted by severe winters and ice build-up due 
to preference for an inshore habitat after the first 
few months of life. An entire population could 
therefore be eliminated by extremes in water 
temperature during winter. In a laboratory study 
to determine if there was a size differential in 



1990 



GOODCHILD: STATUS OF THE BROOK SILVERSIDE 



43 



mortality due to lethal temperature, most deaths 
began when the temperature of the water dropped 
below 7°C. No differential mortality with respect 
to size was observed, however (Nelson 1968). 

Predation by other species may be a limiting 
factor for Brook Silverside because they are 
common forage for many species. Avoidance of 
shallow water by juveniles may protect the 
population from excessive predation. As they are 
for the most part the only inhabitants of these 
waters, they escape the competition existing in 
shallow water. Small size would make them ideal 
food for the dozens of species which feed in the 
shallows along the shore. In studies of bass 
populations in the southern United States, 
Timmons et al. (1980) reported that small 
Largemouth Bass {Micropterus salmoides) 
selected Brook Silverside in May but later avoided 
them and Largemouth Bass larger than 100 mm 
consumed none. 

Special Significance of the Species 

The Brook Silverside is the only member of the 
family Atherinidae that lives in Canadian 
freshwaters. Atherinids compose a family whose 
members are typically salt water forms, none of 
which attains a large size. 

Although widely distributed throughout the 
freshwaters of central North America, Labidesthes 
sicculus is confined to a small region in south- 
central Canada. It is undoubtedly of very minor 
importance in Canadian waters because of its low 
numbers and limited distribution (Scott and 
Crossman 1973). It is, however, an ideal forage fish 
and may be an important prey for game fishes 
when abundant. In the United States, larger 
specimens are occasionaly used for bait by sports 
fishermen but they are very sensitive to low oxygen 
and die very quickly in a bait bucket (Cooper 
1983). They are not legal baitfish species in 
Ontario. 

There is little public interest in the Brook 
Silverside and these tiny fishes are often overlooked 
or considered to be just another minnow. They are 
exceptional in their surface feeding mechanics, in 
the pelagic habitat requirements of juveniles and in 
their one-year life cycle. 

Evaluation 

The Brook Silverside is an ideal forage fish and 
when abundant, may be an important prey species. 
The status of Canadian populations is not fully 
understood. Collection records do not indicate any 
decline in Brook Silverside populations and 
perhaps its range is expanding slightly. Although it 
is not apparently in any immediate jeopardy, 
populations can be totally eliminated in an 
extremely short time span because of its one year 



life cycle. Excessive turbidity and interspecific 
competition are apparently primarily responsible 
for the decline of Brook Silverside in areas of the 
United States. 

The Brook Silverside is indigenous to Canada and 
at the fringe of its range here. However, Canadian 
populations are contiguous with those in the United 
States and the species appears to be reasonably 
abundant and not particularly vulnerable at this 
time. For these reasons classification by COSEWIC 
is not necessary at this time, however its status 
should be re-examined at regular intervals. 

Acknowledgments 

Financial support for the preparation of this 
report was made possible by World Wildlife Fund 
(Canada), the Department of Fisheries and Oceans 
and Environment Canada. Sincere thanks to R. R. 
Campbell, Department of Fisheries and Oceans 
for his support in the preparation of this report. 
Also to W. B. Scott, Huntsman Marine Labora- 
tory, and E. J. Crossman, Royal Ontario Museum, 
for permission to use the drawing of Labidesthes 
sicculus from Freshwater Fishes of Canada. 

The assistance of D. E. McAllister of the National 
Museum of Natural Sciences (now Canadian 
Museum of Nature); E. J. Crossman, E. Holm and 
R. Winterbottom, of the Royal Ontario Museum; 
and G. E. Gale and G. A. Goodchild, of the Ontario 
Ministry of Natural Resources, in providing access 
to records and reports was invaluable. N. E. 
Mandrak, Department of Zoology, University of 
Toronto, Toronto, Ontario, provided valuable 
insight on the distribution of this species. I am 
particularly grateful to A. and D. Herbert for 
logistical support. 

Literature Cited 

Bailey, R. M., and G.R. Smith. 1981. Origin and 

geography of the fish fauna of the Laurentian Great 

Lakes Basin. Canadian Journal of Fisheries and 

Aquatic Science 38: 1539-1561. 
Bailey, R. M., H. E. Winn, and C. L. Smith. 1954. 

Fishes from the Escambia River, Alabama and 

Florida, with ecologic and taxonomic notes. 

Proceedings of the Academy of Natural Science of 

Philadelphia 106: 109-164. 
Bangham, R. V. 1972. A resurvey of the fish parasites of 

Western Lake Erie. Bulletin of the Ohio Biological 

Survey 4(2): 1-23. 
Bangham, R. V., and G. W. Hunter, IIL 1939. Studies 

on fish parasites of Lake Erie. Distribution studies. 

Zoologica 24(4): part 27: 385-448. 
Bensley, B. A. 1915. The fishes of Georgian Bay. 

Contribution of the Canadian Biological Society 

Sessional Paper 39b (191 1-1914): 1 51. 
Berg, R. E., P. A. Doepke, and P. R. Hannuksela. 1975. 

First occurrence of the Brook Silverside, Labidesthes 

sicculus, in a tributary of Lake Superior. Journal of the 

Fisheries Research Board of Canada 32: 2541 2542. 



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The Canadian Field-Naturalist 



Vol. 104 



Boesel, M. W. 1938. Food of nine species of fish from 
the western end of L. Erie. Transactions of the 
American Fisheries Society 67: 215-223. 

Cahn, A. R. 1927. An ecological study of southern 
Wisconsin fishes: the Brook Silverside {Labidesthes 
sicculus) and the cisco {Leucichthys artedi) in their 
relations to the region. lUinois Biological Monographs 
11(1): 1 151. 

Clay, W. M. 1962. A field manual of Kentucky fishes. 
Kentucky Department of Fish and Wildlife Resources. 
147 pages. 

Cooper, E. L. 1983. Fishes of Pennsylvania and the 
northeastern United States. Pennsylvania State 
University Press, University Park, Pennsylvania. 243 
pages. 

Cross, F. B. 1967. Handbook of fishes of Kansas. 
University of Kansas Museum of Natural History, 
Miscellaneous Publication 45. 357 pages. 

Denoncourt, R. P., E. C. Raney, G. H. Hocutt, and J. R. 
Stauffer. 1975. A checklist of the fishes of West 
Virginia. Virginia Journal of Science 26(3): 1 17-120. 

Fogle, N. E. 1959. Someaspectsof the life history of the 
brook silversides, Labidesthes sicculus, in Lake Fort 
Smith, Arkansas. M.Sc. thesis, University of 
Arkansas, Little Rock, Arkansas. 25 pages. 

Giesy, J. P., Jr., and J. G. Wiener. 1977. Frequency 
distributions of trace metal concentrations in five 
freshwater fishes. Transactions of the American 
Fisheries Society 106(4): 393-403. 

Harlan, J. R., and E. B. Speaker. 1956. Iowa fish and 
fishing. Iowa State Conservation Commission. 377 
pages. 

Hoffman, G. L. 1967. Parasites of North American 
freshwater fishes. University of California Press, Los 
Angeles, California. 486 pages. 

Hubbs, C. L. 1921. An ecological study of the life- 
history of the fresh-water atherine fish Labidesthes 
sicculus. Ecology 2(4): 262-276. 

Hubbs, C. L., and K. F. Lagler. 1967. Fishes of the 
Great Lakes Region. University of Michigan Press, 
Ann Arbor, Michigan. 213 pages. 

Johnson, J. E. 1987. Protected fishes of the United 
States and Canada. American Fisheries Society, 
Bethseda, Maryland. 42 pages. 

Keast, A., and D. Webb. 1966. Mouth and body form 
relative to breeding ecology in the fish fauna of a small 
lake. Lake Opinicon, Ontario. Journal of the Fisheries 
Research Board of Canada 23(12): 1845-1867. 

Keast, A., and J. Eadie. 1984. Growth in the first 
summer of life: a comparison of nine co-occurring fish 
species. Canadian Journal of Zoology 62: 1242 1250. 

Korth, J. W., and J. M. Fitzsimmons. 1987. Karyology 
of three species of eastern North American atherinid 
fishes. Copeia(2): 505 509. 

Leach, J. H., and S.J. Nepszy. 1976. The fish 
community in Lake Erie. Journal of the Fisheries 
Research Board of Canada 33: 622 638. 

Lee, D. S. 1980. %roo]f.':i\\vcys\dQ, Labidesthes sicculus. 
Page 557 in Atlas of North American freshwater fishes. 
Edited by D. S. Lee,C. R. Gilbert, C. H. Hocutt, R. E. 
Jenkins, D. E. McAllister, and J. R. Stauffer, Jr. 
North Carolina Stale Museum of Natural History 
Biological Survey Publication 1980 12. 



McAllister, D. E., and B. W. Coad. 1974. Fishes of 

Canada's National Capital Region. National Museum 

of Natural Sciences Miscellaneous Special Publication 

24. 200 pages. 
McComas, S. R., and R. W. Drenner. 1982. Species 

replacement in a reservoir fish community; silverside 

feeding mechanics and competition. Canadian Journal 

of Fisheries and Aquatic Science 39: 815-821. 
Mandrak, N. E. 1990. The zoogeography of Ontario 

freshwater fishes. M.Sc. thesis. University of Toronto, 

Toronto, Ontario. 
Miller, R. R. 1972. Threatened freshwater fishes of the 

United States. Transactions of the American Fisheries 

Society 101(2): 239-252. 
Nelson, J. S. 1968. Life history of the Brook Silverside, 

Labidesthes sicculus, in Crooked Lake, Indiana. 

Transactions of the American Fisheries Society 97: 

293-296. 
Pflieger, W. L. 1975. The fishes of Missouri. Missouri 

Department of Conservation. 343 pages. 
Radforth, I. 1944. Some considerations of the distribu- 
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Zoology Contribution 25: 1-116. 
Rasmussen, R. P. 1980. Egg and larva development of 

Brook Silversides from the Peace River, Florida. 

Transactions of the American Fisheries Society 109: 

407-416. 
Schmidt, R. E. 1986. Zoogeography of the northern 

Appalachians. Pages 137-160 /«The Zoogeography of 

North American freshwater fishes. Edited by C. H. 

Hocutt and B. O. Wiley. John Wiley and Sons, New 

York. 866 pages. 
Scott, W. B. 1967. Freshwater fishes of eastern Canada. 

Second edition. University of Toronto Press, Toronto. 

137 pages. 
Scott, W. B., and M. G. Scott. 1988. Atlantic fishes of 

Canada. Canadian Bulletin of Fisheries and Aquatic 

Sciences Number 219. 
Scott, W. B., and E.J. Grossman. 1973. Freshwater 

fishes of Canada. Fisheries Research Board of Canada 

Bulletin 184: 1 966. 
Smith, C.L. 1985. Theinlandfishesof New York State. 

New York State Department of Environmental 

Conservation, Ithaca, New York. 522 pages. 
Smith, P. W. 1979. The fishes of Illinois. University of 

Illinios Press, Chicago, Illinois. 314 pages. 
Smith- Vaniz, W. F. 1968. Freshwater fishes of Ala- 
bama. Auburn University, Agricultural Experiment 

Station, Auburn, Albama. 21 1 pages. 
Timmons, T. J., W. L. Shelton, and W. D. Davies. 

1980. Differential growth of largemouth bass in West 

Point Reservoir, Alabama-Georgia. Transactions of 

the American Fisheries Society 109: 176-186. 
Toner, G. C. 1943. Ecological and geographical 

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Toronto, M.A. thesis, Toronto, Ontario. 91 pages. 
Trautman, M. B. 1981. The fishes of Ohio with 

illustrated keys. Revised edition. Ohio State University 

Press, Columbus, Ohio. 782 pages. 

Accepted 10 October 1989 



Status of the Banded Killifish, Fundulus diaphanus, in Canada* 
J. Houston 

374 Fireside Drive, Woodlawn, Ontario KOA 3M0 

Houston, J. 1990. Status of the Banded Killifish, Fundulus diaphanus, in Canada. Canadian Field-Naturalist 104(1): 
45-52. 

The Banded Killifish, Fundulus diaphanus, is one of three species of the genus occurring in Canadian waters and is a 
species preferring the quiet waters of lakes and ponds. It tolerates temperatures in excess of 30° C, but prefers 
temperatures in the range of 21° to 28° C. They are often used by anglers as bait fish and can survive in water with low 
oxygen content for extended periods of time. In Canada, the Banded Killifish is widely distributed in the Maritime 
Provinces but is known from only a few localities in Newfoundland. The range extends west through southern Quebec 
and Ontario in the Great Lakes watershed to Lake Huron. They have been recorded from the Red River at Winnipeg, 
Manitoba, Crowduck Lake, Manitoba and Lake of the Woods, Ontario. The species has an extremely limited 
distribution in Newfoundland and Manitoba where further immigration in these areas is unlikely due to lack of 
suitable habitat or barriers to dispersion. 

Le fondule barre, Fundulus diaphanus, I'une des trois especes du genre Fundulus que Ton trouve au Canada, prefere 
les eaux calmes des lacs et des etangs. Ce poisson tres robuste tolere des temperatures depassant 30° C bien qu'il soit 
plus a I'aise dans des eaux ou la temperature se situe entre 2 1 ° et 28° C. Les pecheurs a la ligne I'utilisent souvent comme 
appat, et il est capable de survivre dans une eau tres pauvre en oxygene pendant une periode prolongee. Au Canada, le 
fondule barre est largement repandu dans les Maritimes, sauf a Terre-Neuve ou il n'a ete observe qu'a quelques 
endroits seulement. Son aire de repartition s'etend vers I'ouest dans le sud du Quebec et de I'Ontario, et le bassin des 
grands Lacs jusqu'au lac Huron. On a signale sa presence a deux endroits au Manitoba, soit dans la riviere Rouge a 
Winnipeg et le lac Crowduck, ainsi que dans le lac des bois en Ontario. Le fondule barre a une repartition extremement 
limite a Terre-Neuve et au Manitoba ou I'implantation de I'espece parait improbable a cause de barrieres qui 
empechent sa dispersion ou de I'absence d'habitats appropries. 

Key Words: Banded Killifish, Fundulus diaphanus, fondule barre, Fundulidae, killifishes, topminnow, rare and 
endangered fishes, status, Canada. 



The Banded Killifish, Fundulus diaphanus 
(Lesueur 1817), is a member of the killifish or 
topminnow family (Fundulidae). These are small 
fishes with a wide distribution in the fresh and salt 
waters of North and Central America. They occur 
also in Europe, Africa, Asia and the East Indies; 
they are most common in the southeast United 
States (Scott and Grossman 1973). The genus 
Fundulus, of the 45 nominal genera, is represented 
in Canada by three species: Fundulus diaphanus, 
Fundulus heteroclitus, and Fundulus no tat us. 
These fishes are often referred to as topminnows 
because of their habit of surface feeding (Leim and 
Scott 1966; Scott and Crossman 1973). 

Banded Killifish (Figure 1) average 6 to 8 cm in 
total length (TL), with a maximum known size of 
11.4 cm (Scott and Crossman 1973). Like most 
topminriows, the mouth is small and directed 
upwards. The lower jaw protrudes beyond the 
upper when the mouth is closed. These fish have an 
overall brown to olive-green colouration dorsally, 
shading to white or yellow below. There are many 
(12 to 20) dark, vertical bars on the sides, more 



widely spaced in females, and these may be 
accentuated during spawning. The vertical bars 
and overall colouration are more intense during 
spawning and the dorsal fin may show a greenish 
or golden wash as well as the hint of a dark band or 
two (Scott and Crossman 1973). 

The Blackstripe Topminnow {Fundulus nota- 
tus) is known in Canada only from the Sydenham 
River system in southwestern Ontario {see 
McAllister 1987), whereas the Mummichog 
{Fundulus heteroclitus) is restricted to the brackish 
shore waters of the Atlantic provinces, including 
southwestern Newfoundland, and to the estuaries 
and salt marshes of the Gulf of St. Lawrence to the 
head of the tide above Anticosti Island (Scott and 
Grossman 1973). The Banded Killifish is widely 
distributed in the Maritime Provinces, Quebec, 
and Ontario; its limited distribution in Manitoba 
and Newfoundland begs an examination of its 
status in Canada. This report documents the 
current status of the species in Canada for the 
Committee on the Status of Endangered Wildlife 
in Canada (COSEWIC). 



* Vulnerable status approved and assigned by COSEWIC 11 April 1989. 

45 



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The Canadian Field-Naturalist 



Vol. 104 




Figure I. Drawing of the Banded Killifish, Fundulus diaphanus [adapted from Scott 
and Grossman (1973) by permission]. 



Distribution 

Banded Killifish occur along the eastern 
seaboard from South Carolina north to the 
Maritimes and west through southern Canada 
(Figure 2) to the Red River of Manitoba and the 
Yellowstone River of eastern Montana in the 
United States (Scott and Crossman 1973; Gilbert 
and Shute 1980). 

In Canada, the species is widely distributed 
(Figures 2,3) in the Atlantic Provinces, with the 
exception of Newfoundland where it is known only 
from a few localities in the southwest and from the 
Burin Peninsula (Scott and Crossman 1973; 
Gibson et al. 1984). The Canadian distribution 
continues west through Quebec in the St. 
Lawrence valley where suitable habitat may be 
found and on into the Great Lakes watershed of 
southern Ontario. The species is not known from 
Lake Superior (Scott and Crossman 1973), but has 
been recorded from Lake of the Woods (Stewart et 
al. 1985). In Manitoba, the species has only been 
recorded from the Red River at Winnipeg and 
Crowduck Lake (Winnipeg River Watershed), 
near the Ontario border (Stewart et al. 1985). 

Protection 

No specific protection for the species exists in 
Canada. General protection is available through 
the habitat sections of the Fisheries Act. The 
species is considered to be of special concern in the 
Province of Manitoba (Johnson 1987). In Ontario, 
the species is not classed as a bait fish and it is 
illegal to use it as such. 

In the United States, Banded Killifish are 
considered to be endangered in Pennsylvania and 
South Dakota (Miller 1972), and disappearing 
from Illinois (Smith 1979). The species has 
protected status in the states of Ohio and South 
Dakota due to its rarity (Johnson 1987). 

Population Si7.es and Trends 

As for most small fishes, such as the minnows, 
no population estimates have been made for the 
topminnows. In many parts of the range there is no 
direct evidence for trends in population size. 



The species is locally abundant in parts of its 
United States range, but has undergone a decline 
since the 1920s (Trautman 1957). This decline is 
particularly evident in the western sectors of the 
distribution where the species remains in a few 
lakes of glacial origin in Illinois (Smith 1979) and 
has disappeared from many localities in Ohio 
(Trautman 1957) and is considered to be 
endangered in Pennsylvania and South Dakota 
(Miller 1972; Johnson 1987). 

Within the known Canadian range (Figure 3), 
Banded Killifish appear in moderate abundance 
and collection records indicate that the range may 
be stable [National Museum of Natural Sciences 




Figure 2. Approximate North American range of the 
Banded Killifish, Fundulus diaphanus. 



1990 



HOUSTON: Status of the Banded Killifish 



47 




Figure 3. General distribution of Canadian collection records of the Banded Killifish, Fundulus 
diaphanus. Closed circles represent collections made prior to 1980, triangles represent post 1980 
collections. (Based on sources referenced in the text. Some collections have been omitted due to map 
scale). 



(NMC), now Canadian Museum of Nature; Royal 
Ontario Museum (ROM); Ontario Ministry of 
Natural Resources (OMNR) records]. The species 
is particularly abundant in Nova Scotia (Fritz and 
Garside 1977). 

In insular Newfoundland, the species has been 
recorded from the southwest corner of the island 
near Stephenville Crossing at the head of St. 
George's Bay (48° N, 59° W) and in the Highlands 
River 50 km south of Stephenville (Scott and 
Grossman 1964; Gibson et al. 1984). In 1983, 
Banded Killifish were collected from Freshwater 
Pond on the Burin Peninsula (47°6'N, 55°16'W); 
this is the most easterly occurrence recorded for the 
species (Gibson et al. 1984). The species seems to be 
abundant at these locations and breeding 
populations appear to exist; however, suitable 
habitat along the coast is limited, and the steep 



gradient of the rivers may constitute barriers to 
immigration to sites further inland (Gibson et al. 
1984). 

Species records [NMC; ROM; OMNR] are 
sparse along the North Shore of the Gulf of St. 
Lawrence and the St. Lawrence River to Quebec. 
The most northerly distribution reported for the 
species is at the Matamek River (50°18'N, 
65°57'W) on the North Shore (Gibson and Sears 
1977) where the fish were relatively abundant. 
Such occurrences should be expected due to the 
proximity of other collections. However, as in 
insular Newfoundland, this coast is rocky and 
suitable habitat is probably limited. 

In Ontario and Quebec, the distribution and 
abundance would appear to be relatively stable as 
recent collection records confirm the presence of 
the species in the same watersheds where collected 



48 



The Canadian Field-Naturalist 



Vol. 104 



earlier in the century (e.g. Mongeau et al. 1972; 
McAllister and Coad 1974; NMC, OMNR, and 
ROM records). With the exception of recent 
collections from Lake of the Woods (see Stewart et 
al. 1985), no range extensions have been reported. 
The presence of the species in Shoal Lake in the 
Lake of the Woods region should not be surprising 
as Stewart et al. (1985) also report recent (1982) 
evidence for the species in Crowduck Lake, 
Manitoba (50°05'N, 95°08'W), part of the same 
Winnipeg River drainage system. 

Banded Killifish are known in Manitoba only 
from one other collection from the Red River at 
Winnipeg (Stewart-Hay 1954), although the 
species is known from the Red River drainage in 
North Dakota and Minnesota (Eddy and 
Underbill 1974; Owen et al. 1981). Stewart et al. 
(1985) considered the species to be extremely rare 
in Manitoba where it has not otherwise been found 
despite frequent collecting efforts. 

Habitat 

Scott and Grossman (1973) describe suitable 
habitat as quiet waters of lakes and ponds with 
sand, gravel, or detritus-covered bottoms and 
patches of submerged aquatic vegetation. The 
species is most abundant in very shallow waters 
and has a preference for clear, glacial lakes with 
sluggish waters and abundant vegetation (Traut- 
man 1957; Smith 1979; Cooper 1983). Water 
clarity is important as the species relies on visual 
perception in prey selection (Desgagne and 
Lalancette 1984). 

Banded Killifish appear to be tolerant of low 
dissolved oxygen. Although O2 limits have 
apparently not been determined, these fish have 
been known to exist for long periods of time in 
minnow buckets and even out of water (see Scott 
and Grossman 1973). They also have a wide 
tolerance of water temperatures and have been 
known to exist at temperatures to 38.3°G 
(Garlander 1969), ahhough Rombough and Garside 
(1977) reported an upper incipient limit of 34.5° G. 
Melisky et al. (1980) indicated that the distribution 
within a water system may be influenced by 
temperature and found that fish in Pennsylvania 
have a preference for waters in the neighborhood of 
28.6°G while those in Nova Scotia exhibit a 
preference of 21.0°G. In Quebec, prespawning 
activities became apparent at 21°G and spawning 
probably occurs when water temperature reaches 
23° G (Scott and Grossman 1973). 

The species is euryhaline, but usually inhabits 
freshwater streams and lakes, rarely being found in 
brackish or marine waters (Fritz and Garside 
1974b, 1975). in Nova Scotia, they are abundant in 



certain brackish lakes where their fecundity 
appears to be higher than in nearby oligotrophic 
lakes possibly because of the greater productivity 
in the former (Fitz and Garside 1975). Although 
the species has an ultimate salinity preference for 
freshwater, they acclimate easily to salt water 
(Fritz and Garside 1974a; Garside and Morrison 
1977; Weisberg 1986) and can tolerate saUnities in 
excess of 20 ppt (Griffith 1974; Weisberg 1986). 
Griffith (1974) suggested that the apparent 
immediate preference for fresh water was 
acclimation dependent which may explain their 
presence in Prince Edward Island, Anticosti 
Island, and Newfoundland. 

General Biology 

Systematic Notes: Two subspecies of Banded 
Killifish are recognized and both exist in Ganada 
(Scott and Grossman 1973; Robins et al. 1980). 
The Eastern Banded Killifish, Fundulus diaphanus 
diaphanus, is found in the Atlantic drainage and 
the range extends west to the upper St. Lawrence 
River and eastern Lake Ontario. The Western 
form, Fundulus diaphanus menona, ranges 
eastward from the Yellowstone River in Montana 
to Lake Erie. The subspecies integrate along and in 
Lake Ontario and the upper St. Lawrence River 
(Scott and Grossman 1973; Gilbert and Shute 
1980). Hybridization with other species of the same 
genera is said to be rare (Scott and Grossman 
1973), although Banded Killifish do appear to 
readily hybridize with Mummichogs where the 
species are sympatric (Fritz and Garside 1974a). 

Life History: Individuals reach maturity at an 
age of 1+ years and a total length of about 6 cm 
(Garlander 1969). Adults attain maximum size in 
three years (Gooper 1983), but information on 
longevity in the species is lacking. The size of adults 
varies with location, but usually ranges from 6 to 8 
cm. Larger individuals are more common in the 
Maritime Provinces where individuals up to 11.4 
cm in length have been recorded (Scott and 
Grossman 1973). Fritz and Garside (1975) found 
that Banded Killifish from brackish lakes grew 
faster in the first year, attained greater overall size 
and were more fecund than fish from oligotrophic 
lakes. They attributed this difference to the greater 
productivity in the brackish lake, but despite the 
advantage in brackish waters, the species showed a 
preference for fresh water and they concluded that 
salinity for some reason is important in the 
distribution of the species (Fritz and Garside 
1974b, 1975). 

Spawning has been described by Richardson 
(1939) and takes place from April to May 
depending on water temperature, a temperature of 



1990 



HOUSTON: Status of the Banded Killifish 



49 



around 21° C being preferred (Carlander 1969). 
McAllister and Coad (1974) reported finding 
females in spawning condition in the Ottawa River 
watershed in June when water temperature was 
around 23° C. Males select breeding sites in the 
quiet shallows of weedy areas. The males defend 
these territories and develop more intensive 
colouration while the females remain pale (Scott 
and Grossman 1973). Males spawn individually 
with females and there is no nest building or care of 
the young (Cooper 1983). The rriales and females 
seem to pair off according to size and after pursuit 
by the male, the female extrudes her eggs in clusters 
of five to 10 eggs. The eggs are attached to the 
female by a thread and following fertilization, 
detach and stick to vegetation by individual, 
adhesive threads. This continues until all eggs of 
the female are laid {see Scott and Grossman 1973). 

A female may contain up to 250 or so eggs 
(Carlander 1969) of about 2 mm in diameter. The 
eggs hatch in 1 1 to 1 2 days at water temperatures of 
22 to 27° C (Cooper 1936). The fry are 6 to 7 mm 
long and larvae 7 to 12 mm have been described by 
Fish (1932) and Auer (1982). Fritz and Garside 
(1975) reported on growth rates, length-weight 
relationships and fecundity between two popula- 
tions in Nova Scotia and determined that fish from 
brackish waters did better than their counterparts 
in fresh water. They attributed this to the higher 
productivity of the brackish lake. Growth in the 
first year appears to be rapid. From a 6 to 7 mm fry, 
young fish may attain lengths of 2.0 to 6.4 cm by 
the end of the year (Trautman 1957; Smith 1952). 
The largest specimen on record in Canada 
measured 11.4 cm total length and was taken in 
Lake O'Law, Nova Scotia (Scott and Grossman 
1973). 

Parasites of the species have been hsted by 
Bangham and Hunter (1939) and Hoffman (1967). 
Banded Killifish serve as a forage species to game 
fish where the ranges overlap and they occur in 
numbers. It may also be an important item in the 
diet of fish-eating birds such as the American 
Merganser {Mergus merganser) or the Belted 
Kingfisher (Megaceryle alcyon) in some areas 
(White 1943, 1957). 

Behaviour: Spawning behaviour is similar to 
that of other killifishes with males defending 
territories and spawning individually with females 
(Cooper 1983) and has been described in detail by 
Richardson (1939). The species is usually found in 
schools which tend to favour sand-bottom 
shallows (Scott and Grossman 1973; Smith 1979). 
The schooling behaviour may be related to 
predator avoidance {see Godin and Morgan 1985). 
There is no information on seasonal migrations 
but the distrtibution of the fish within a body of 
water is probably related to salinity and 



temperature preferences and to the availability of 
suitable prey (Fritz and Garside 1974a; Griffith 
1974; Garside and Morrison 1977; Weisberg 1986). 

Food and Feeding: Banded Killifish are versatile 
feeders and despite the superior position of the 
mouth, they utilize all levels of the water column 
(Keast and Webb 1966). Smaller individuals (less 
than 4 cm) eat chironomid larvae, ostracods, 
cladocerans, copepods, and small quantities of 
amphipods and even flying insects. Larger 
individuals take the same items, but also are 
known to eat Odonata and Ephemeroptera 
nymphs, molluscs, tubellarians, and small 
crustaceans (Keast and Webb 1966; Scott and 
Grossman 1973; Baker-Dittus 1978). 

Prey selection relies on visual perception 
(Desgagne and Lalancette 1984) and feeding is as a 
member of a school (Keast and Webb 1966). 
Competition with Fundulus hereroclitus, where 
the two are sympatric, seems to be minimized by 
differences in diet and foraging patterns (Baker- 
Dittus 1978; Weisberg 1986). 

Limiting Factors 

The availability of suitable habitat and barriers 
to access may be limiting to the distribution of the 
species in Newfoundland and along the North 
Shore of the Gulf of St. Lawrence (Gibson and 
Sears 1977; Gibson etal. 1984). The lack of suitable 
habitat [clear lakes of glacial origin with much 
aquatic vegetation (Trautman 1957; Smith 1979) 
could be limiting further distribution in Manitoba 
as well. Clear water with abundant vegetation 
seems necessary to the survival of the species 
(Trautman 1957). Clear water may be important 
due to the reliance of visual perception in prey 
selection (Desgagne and Lalancette 1984) and 
vegetation may be important for predator 
avoidance (Godin and Morgan 1985), foraging 
patterns (Baker-Dittus 1978) and reproduction 
(Richardson 1939). Notwithstanding, there are still 
many lakes in both Manitoba and Newfoundland 
which would qualify as suitable habitat and which 
are also accessible to known collection sites, even 
to small fish with limited ability to ascend rapids 
etc. (J. Gibson, Department of Fisheries and 
Oceans, St. John's, Newfoundland; K. W. 
Stewart, Department of Zoology, University of 
Manitoba, Winnipeg, Manitoba; personal com- 
munications). Both K. W. Stewart and J. Gibson 
(personal communications) suggest that the 
species may be limited by temperature, i.e., the 
cooler waters of Manitoba and Newfoundland are 
colder than the 21° to 23° C necessary for killifish 
production. If this is the case, one might expect an 
increase in abundance and distribution if the recent 
warming trend continues. Other species (e.g. 
Micropterus dolomieui, Morone chrysops. 



50 



The Canadian Field-Naturalist 



Vol. 104 



Poxomis nigromaculatus, N o turns flavus, Notro- 
pis spilopterus have demonstrated this trend in the 
last five to 10 years (K. W. Stewart, personal 
communication). 

There are no indications of decline in Canadian 
populations; however the species has disappeared 
from significant portions of its former range in 
Ohio and Illinois, where it is thought that the 
destruction and general deterioration of natural 
lakes has led to this decline (Trautman 1957; Smith 
1979). Although similar habitat disruption and 
deterioration has occurred in Ontario, for 
example, there is no evidence of a similar decrease 
in populations here, but there have been no 
serious, specific surveys to that end and it may well 
be that there have been losses in certain areas. 

It is interesting to note that it seems to be the 
western form of the species that has shown 
evidence of decline in parts of the United States 
range. In Ohio, both forms are present, but it is 
apparently only the western form that has been 
affected (Trautman 1957). Fundulus diaphanus 
menona exists only from Illinois westward, and 
throughout the western part of the United States 
range, this subspecies seems generally to be in 
trouble (Miller 1972; Johnson 1987), perhaps due 
to increases in turbidity. Temperature and salinity 
preferences may bear some influence on the 
distribution of the species (Fritz and Garside 
1974b; Garside and Morrison 1977; Melisky et al. 
1980; Weisberg 1986) and Fundulus diaphanus 
diaphanus can tolerate brackish waters and 
apparently does well in them (Fritz and Garside 
1975), but for some reason not yet determined, it 
shows a distinct preference for fresh water (Keast 
and Webb 1966; Griffith 1974; Fritz and Garside 
1975). This might be related to competition with 
Fundulus heteroclitus. It may be that the eastern 
form has less stringent requirements for water 
quality and vegetation tha the western form and 
has not been affected to the same degree by habitat 
perturbations. 

Competition with similar and/ or related species 
for food or a niche does not seem to be limiting 
(Fritz and Garside 1975; Baker-Dittus 1978; 
Weisberg 1986), although Labidesthes sicculus is 
also a surface feeder. Predation by game fish, 
waterfowl or other fish-eating birds could limit 
populations in some areas (White 1953, 1957; Scott 
and Grossman 1973). 

Chronic exposure to aquatic contaminants may 
produce physiological symptoms and death offish. 
Studies of the effects of first generation herbicide 
and pesticide reagents have shown that many of 
these had many undesirable side effects and most 
have been withdrawn from use, although residues 
of some are persistent and will remain in the 
environment for some time (Rehwoldt et al. 1977). 



Second generation reagents commonly in use are 
more specific and shorter lived. Experiments 
carried out with Banded Killifish from the Hudson 
River indicate that there were no substantial effects 
to the species from chronic exposure to chlorinated 
hydrocarbons or organic phosphates at levels 
present in the river water (Rehwoldt et al. 1977). 

Special Significance of the Species 

Banded Killifish are interesting and colourful 
fish that do well in aquaria and may be useful to 
science in toxicology experiments, etc. They may 
be an important forage species to game fish in some 
areas and provide food for fish-eating birds as well. 
They are of little or no economic importance, but 
are sometimes used as bait fish in the Maritime 
Provinces. Absence of the species from former 
range may be indicative of changing water quality. 

Evaluation 

It appears that the species is widely distributed in 
Canada and, in most areas, is locally abundant. 
The species is known from only two locations in 
Manitoba where it is not abundant and has not 
been found elsewhere in the province despite 
various collection attempts. The species is 
extremely rare there, but contiguous with 
populations in Ontario, North Dakota, and 
Minnesota. Likewise in Newfoundland, the species 
has only been recorded from two widely separated 
localities, but appears to be more locally abundant 
than in Manitoba. It is unlikely that the 
distribution here is much broader than that already 
reported because of barriers (physical and 
climatic) to further immigration. Until further 
evidence proving a wider distribution and 
abundance for the species in Newfoundland is 
available, it should be considered rare and 
vulnerable in that province as these populations 
are discrete from other Canadian populations. 

Acknowledgments 

The author is grateful to the Committee on the 
Status of Endangered Wildlife in Canada 
(COSEWIC) for the opportunity to prepare the 
report. Financial support was made available 
through cooperative funding arrangements 
between The Department of Fisheries and Oceans, 
Environment Canada, and World Wildlife Fund 
Canada. I would also like to acknowledge the 
advice and assistance of Don McAllister of the 
National Museum of Natural Sciences (now 
Canadian Museum of Nature) and for access to 
museum and literature records. The Royal Ontario 
Museum and the Ontario Ministry of Natural 
Resources are also acknowledged for provision of 
collection record information. 



1990 



HOUSTON: Status of the Banded Killifish 



51 



Literature Cited 

Auer, N. A. Editor. 1 982. Identification of larval fishes of 
the Great Lakes Basin with emphasis on the Lake 
Michigan Drainage. Great Lakes Fisheries Commission 
Special Publication 82 3. 

Baker-Dittus, A. M. 1978. Foraging patterns of three 
sympatric killifish. Copeia 1978(3): 383-389. 

Bangham, R. V., and G. W. Hunter. 1939. Studies on 
fish parasites of Lake Erie. Distribution studies. 
Zoologica 24(4); part 27: 385-448. 

Carlander, K. D. 1969. Handbook of freshwater fishery 
biology, Volume one. Iowa State University Press, 
Ames, Iowa. 

Cooper, E. L. 1983. Fishes of Pennsylvania and the 
northeastern United States. Pennsylvania State 
University Press, University Park, Pennsylvania. 

Cooper, G. P. 1936. Importance of forage fishes. Pages 
305-30 in Proceedings of the First North American 
Wildlife Conference. 

Desgagne L., and L-M. Lalancette. 1984. Role de la 
perception visuelle dans la selectivite des Daphnia 
magna et des Cyclops scutifer chez le fondule barre, 
Fundulus diaphanus. Science et Technique de I'Eau 
17(3): 287-288. 

Eddy, S., and J. C. Underbill. 1974. Northern fishes with 
special reference to the upper Mississippi valley. Third 
edition. University of Minnesota Press, Minneapolis, 
Minnesota. 

Fritz, E. S., and E. T. Garside. 1974a. Identification and 
description of hybrids of Fundulus heteroclitus and F. 
diaphanus (Pisces: Cyprinodontidae) from Porters 
Lake, Nova Scotia, with evidence for absence of 
backcrossing. Canadian Journal of Zoology 52(12): 
1433-1442. 

Fritz, E. S., and E. T. Garside. 1974b. Salinity preferen- 
ces of Fundulus heteroclitus and F. diaphanus (Pisces: 
Cyprinodontidae): their role in geographic distribution. 
Canadian Journal of Zoology 52(8): 997-1003. 

Fritz, E. S., and E. T. Garside. 1975. Comparison of age 
composition, growth, and fecundity between two 
populations each of Fundulus heteroclitus and F. 
diaphanus (Pisces: Cyprinodontidae). Canadian 
Journal of Zoology 53(4): 361-369. 

Garside, E. T., and G. C. Morrison. 1977. Thermal 
preferences of mummichog, Fundulus heteroclitus L., 
and banded killifish, F. diaphanus (LeSeueur), 
(Cyprinodontidae) in relation to thermal acclimation 
and salinity. Canadian Journal of Zoology 55(7): 
1190-1194. 

Gibson, R. J., and R. Sears. 1977. An occurrence of 
Fundulus diaphanus (LeSueur) on the North Shore of 
the Gulf of St. Lawrence. Le Naturahste canadien 104: 
273-274. 

Gibson, R. J., J-P. Thonney, and K. Hillier. 1984. An 
easterly extension in the known range for Fundulus 
diaphanus in Newfoundland. Le Naturaliste canadien 
111:213-214. 

Gilbert C. R., and J. R. Shute. 1980. Fundulus diapha- 
nus (LeSueur) Banded Killifish. Page 513 in Atlas of 
North American freshwater fishes. Edited by D. S. Lee, 
C. R. Gilbert, C. H. Hocutt, R. Awe Jenkins, D. E. 
McAlhster, and J. R. Stauffer, Jr. North CaroUna State 
Museum of Natural History, North Carolina Biological 
Survey Publication Number 1980-12. 



Godin, J-G., and M.J. Morgan. 1985. Predator 
avoidance and school size in cyprinodontid fish, the 
banded killifish (Fundulus diaphanus LeSueur). 
Behavioural Ecology and Sociobiology 16: 105-110. 

Griffitb, R. W. 1974. Environment and saUnity tolerance 
in the genus Fundulus. Copeia 1974(3): 319-331. 

Hoffman, G. L. 1967. Parasites of North American 
freshwater fishes. University of California Press, Los 
Angeles, California. 

Johnson, J. E. 1987. Protected fishes of the United States 
and Canada. American Fisheries Society, Bethesda, 
Maryland. 

Keast, A., and D. Webb. 1966. Mouth and body form 
relative to feeding ecology in the fish fauna of a small 
lake, Lake Opinicon, Ontario. Journal of the Fisheries 
Research Board of Canada 23(12): 1845-1867. 

Leim, A. H., and W. B. Scott. 1966. Fishes of the 
Atlantic coast of Canada. Fisheries Research Board of 
Canada Bulletin Number 155. 

McAllister, D. E. 1987. Status of the Blackstripe 
Topminnow, Fundulus notatus, in Canada. Canadian 
Field-Naturalist 101(2): 219-225. 

McAllister, D. E., and B. W. Coad. 1974. Fishes of 
Canada's National Capital Region. Fisheries Research 
Board of Canada Miscellaneous Special Publication 24. 

Melisky, E. L., J. R. Stauffer, Jr., and C. H. Hocutt. 
1980. Temperature preference of banded killifish, 
Fundulus diaphanus, from southwestern Pennsylvania. 
Copeia 1980(2): 346-349. 

Miller, R. R. 1972. Threatened freshwater fishes of the 
United States. Transactions of the American Fisheries 
Society 101(2): 239-252. 

Mongeau, J-R., A. Courtmanche, G. Masse, and B. 
Vincent. 1972. Cartes de repartition geographique des 
especes de poissons au sud du Quebec, d'apres les 
inventaires ichthyologiques effectues de 1963 a 1972. 
Faune du Quebec Rapport Special Numero 4. 

Owen, B. D., D. S. Elsen, and G. W. RusseU. 1981. 
Distribution of fishes in North and South Dakota basins 
affected by the Garrison Diversion Unit. Fisheries 
Research Unit, University of North Dakota, Grand 
Forks, North Dakota. 

Rehwoldt, R. E., E. Kelley, and M. Mahoney. 1977. 
Investigations into the acute toxicity and some chronic 
effects of selected herbicides and pesticides on several 
fresh water fish species. Bulletin of Environmental 
Contamination and Toxicology 18(3): 361-365. 

Richardson, L. R. 1939. The spawning behaviour of 
Fundulus diaphanus (LeSueur). Copeia 1939(3): 
165-167. 

Robins, C. R., Chairman, R. M. Bailey, C. E. Bond, J. R. 
Brooker, E. A. Lachner, R. N. Lea, and W. B. 
Scott. 1980. A list of common and scientific names of 
fishes from the United States and Canada. American 
Fisheries Society Special Publication Number 12. 

Rombough, P. J., and E. T. Garside. 1977. Hypoxial 
death inferred from thermally induced injuries at upper 
lethal temperature, in the banded killifish, Fundulus 
diaphanus (LeSueur). Canadian Journal of Zoology 
55(10): 1705-1719. 

Scott, W. B., and E. J. Grossman. 1964. Fishes occurring 
in the fresh waters of insular Newfoundland. Queen's 
Printer, Ottawa, Ontario. 

Scott, W. B., and E. J. Grossman. 1973. Freshwater 
fishes of Canada. Fisheries Research Board of Canada 
Bulletin 184. 



52 



The Canadian Field-Naturalist 



Vol. 104 



Smith, M. W. 1952. Limnology and trout angling in 
Charlotte County Lakes, New Brunswick. Journal of 
the Fisheries Research Board of Canada 8(6): 383^52. 

Smith, P. W. 1979. The fishes of Illinois. University of 
Illinois Press, Urbana, Illinois. 

Stewart, K. W., I. M. Suthers, and K. Leaves- 
ley. 1985. New fish distribution records in Manitoba 
and the role of a man-made interconnection between 
two drainages as an avenue of dispersal. Canadian 
Field-Naturalist 99(3): 317-326. 

Stewart-Hay, R. K. 1954. A killifish in Manitoba. 
Canadian Field-Naturalist 68(2): 94. 

Trautman, M. B. 1957. The fishes of Ohio with 
illustrated keys. Ohio State University Press, 
Columbua, Ohio. 



Weisberg, S. B. 1986. Competition and coexistence 

among four estuarine species of Fundulus. American 

Zoologist 26: 249-257. 
White, H. C. 1953. The eastern belted kingfisher in the 

Maritime Provinces. Fisheries Research Board of 

Canada Bulletin 97. 
White, H. C. 1957. Food and the natural history of 

mergansers on salmon waters in the Maritime Provinces 

of Canada. Fisheries Research Board of Canada 

Bulletin 116. 



Accepted 10 October 1989 



Status of the Least Darter, Etheostoma microperca, in Canada* 
Ken W. Dalton 

8 Suffolk Street, Nepean, Ontario K2G 3P4 

Dalton, Ken, W. 1990. Status of the Least Darter, Etheostoma microperca, in Canada. Canadian Field-Naturalist 
104(1): 53-58. 

Least Darters, Etheostoma microperca, are very small fish found in clear, quiet, well vegetated waters in central North 
America and the Ozark uplands. The Canadian distribution is restricted to southern Ontario although this is not the 
northern fringe of the species range. Both American and Canadian ranges have declined, but remaining Ontario 
populations seem to be stable. These fish do not presently appear to be vulnerable in Canada, but the status should be 
re-evaluated in a few years. Its need for clear water exposes the species to loss of habitat through human activities. 
Directed surveys are necessary for proper evaluation because of the Least Darters' small size and elusive habits. 

Les petits dards, Etheostoma microperca, sont des minuscules poissons de la famille des percides vivant dans les plans 
d'eau limpides et calmes ou la vegeration est dense du centre de I'Amerique du Nord et des hautes terres des monts 
Ozark. Au Canada, ce poisson n'a ete signale que dans le sud de I'Ontario, bien qu'il ne s'agisse pas la de la limite 
septentrionnale de son aire de repartition. Meme si, aux Etats-Unis et au Canada, ses aires de distribution se sont 
retrecies, les populations qui persistent en Ontario semblent stables. Ces poissons ne semblent pas actuellement etre 
vulnerable au Canada, mais il faudra reevaluer leur situation dans quelques annees. L'espece a besoin d'eau claire et la 
degradation de son habitat a cause des activites d'homme I'expose en danger. Des etudes dirigees s'imposent afin 
d'evaluer de fa9on appropriee la situation des petits dards, car ils sont petits et ont tendance a se dissimuler. 

Key Words: Least Darter, Etheostoma microperca, petit dard, darters, Percidae, rare and endangered fishes, status, 
Canada. 



Least Darters, Etheostoma microperca Jordan 
and Gilbert 1887, (Figure 1 ) are among the smallest 
fish, and smallest vertebrates, found in Canada 
(Scott and Grossman 1973). Adults are only 
25 mm in total length (TL), though females are 
larger than males and may reach 38 mm TL (Scott 
and Grossman 1973). Bodies of the fish are robust 
and laterally compressed. They have relatively 
small heads with equally small mouths, but 
moderately-sized to large eyes. Two dorsal fins are 
present, very close together. The most anterior of 
the two has six or seven relatively strong spines 
while the second holds eight or nine soft rays. The 
caudal fin is rounded and clearly barred. The anal 
fin is well developed with one or two spines. Pelvic 
fins are located low on the ventral surface, close 
behind the pectoral fins (Scott and Grossman 
1973). 

The fish are generally brown in colour, with 
darker brown spots on the back and sides. A 
distinct, but short, dorsal stripe is present anterior 
to the dorsal fins, and eight to 10 dark squarish 
blotches on each side. Males are colourful during 
breeding season, with intensely vivid red-orange 
pelvic and anal fins, and orange spots between the 
spines of the first dorsal fin (Scott and Grossman 
1973; Kuehne and Barbour 1983). The only 
breeding colours exhibited by females are a 



yellowish tinge on the pelvic, pectoral, and anal 
fins (Scott and Grossman 1973). 

Breeding males develop sheets of skin on their 
pelvic fins, which extend out, and give a slight 
concavity to these fins (Petravicz 1936). Nuptial 
tubercles are also present on the rays of pelvic fins 
on breeding males, particularly on the distal 
portions (Burr and Page 1979). Together, the two 
pelvic fins form a saddle-like structure used for 
mounting females (Petravicz 1936). Female Least 
Darters have a distinctive, swollen, conical genital 
papilla, close behind the anus, that can be used for 
sexing at all times of the year (Burr 1978). 

Distribution 

The Least Darter occurs in central North 
America, from southern Ontario west to the 
Michigan-Huron basin (Figure 2) and the southern 
tributaries of Lake Superior (Kuehne and Barbour 
1983). The northern fringe of the range is in North 
Dakota and Minnesota. The range extends south 
as far as the Mississippi River system in 
Minnesota, and further south are disjunct 
populations in the Ozark Uplands; from central 
Missouri to southeastern Oklahoma (Kuehne and 
Barbour 1983). 

In Ganada, Least Darters are found only in 
southern Ontario (Figure 3a); in the western 



''Reviewed and accepted by COSEWIC 1 1 April 1989 — no designation required. 

53 



54 



The Canadian Field-Naturalist 



Vol. 104 




1 m m 
Figure 1. Least Darter, Etheostoma microperca, male (adapted from Burr and Page 
1979). 



tributaries of Lake Ontario, the drainage of Lakes 
Erie and St. Clair, and the southeast drainage of 
Lake Huron (Scott and Grossman 1973). 

These small fish are often not detected in surveys 
because of their small size and secretive habits 
(Trautman 1957; Scott and Grossman 1973; Smith 
1979). They may occur in some areas completely 
unknown to investigators until finally being dis- 
covered through lengthy, repeated sampling (Traut- 
man 1 957) or through the use of seines with a smaller 
than standard mesh size (Scott and Grossman 1973). 
A keen eye can detect their darting movements in 
clear water as they dive and swim into the sand. For 
these reasons, Trautman (1957) assumed that Least 
Darters probably had a more extensive, but 
undiscovered, range in the United States prairies 
before 1900, but is no larger present in the area 
because of extensive habitat modifications. The 
same can be true for many Lake Ontario tributaries. 

The remote Ozark populations may have been, 
until recently, connected to the northern groups of 
Etheostoma microperca by an undetected popula- 
tion or series of populations (Kuehne and Barbour 
1983). Pflieger (1971), however, has suggested that 
the Ozark groups are naturally separated from other 
populations of the species because of glaciation and 
geomorphological processes, and that Ozark 
populations are "glacial relicts." 

Protection 

Least Darters receive legal protection in the state 
of Iowa, but not in other states though the fish are of 
special concern in Kansas, Kentucky and Wisconsin 
(.Johnston 1987). 

The species receives no specific protection in 
Canada but is given incidental protection through 
the fish habitat sections of the Fisheries Act. 

Population Sizes and Trends 

No surveys of the Least Darter have taken place, 
thus populations sizes arc difficult to assess. 



However, the presence or absence, and occasionally 
the relative abundance, can be inferred through 
collection records of the Canadian Museum of 
Nature (NMG), the Royal Ontario Museum 
(ROM), the Ontario Ministry of Natural Resources 
(OMNR), and others. 

Only a few collections of Etheostoma microperca 
have ever been taken from Lake Ontario tributaries 
(Figure 3b). The presence (or former presence) has 
been indicated in the Moira River (ROM 24215), 




Figure 2. North American distribution of the Least 
Darter, Etheostoma microperca. 



1990 



Dalton: Status of the Least Darter 



55 




80 78" 76^ 

Figure 3. Collection records of Etheostoma microperca in Ontario; A. post 1970, B. pre-1970. 



56 



The Canadian Field-Naturalist 



Vol. 104 



Duffins Creek (ROM 24454), the outlet from 
Grenadier Pond in Toronto (ROM 08534), and the 
Credit River (ROM 04334). The last of these 
collections was from Duffins Creek in 1965 (ROM 
24454). Surveys carried out in these waters since 
1965 have not yielded any Least Darters (Holm and 
Crossman 1986a, 1986b). The possibility of the 
species continued undetected presence in the region 
is very small because of extensive urbanization in 
much of the area with its accompanying pollution 
and habitat destruction. 

Etheostoma microperca have not been recorded 
from the Sauble River since July 1958 (ROM 
20120). The numbers captured at that time, 15 
specimens, indicates a relatively good abundance. 
Probably, little habitat alteration has occurred in 
this river except near the mouth at Lake Huron. The 
apparent paucity of collections from the river is 
most likely due to a lack of sampling, and not 
because of a reduction in range or numbers of the 
species (R. R. Campbell, Department of Fisheries 
and Oceans, Ottawa, Ontario; personal communi- 
cation). Another river in the same watershed, 
Arkwright Creek, appears to harbour Least Darters 
in at least three locations (OMNR S83). Although 
abundance can not be determined, the collections do 
indicate the continuing presence of Least Darters in 
the watershed. 

There is no evidence for any reductions in range 
or population size elsewhere in Ontario (Figure 3a). 
The NMC, OMNR, ROM and others collected 
extensively in Ontario waters in the early to mid 
1970s. Numbers taken, and the numerous collection 
sites reveal that Least Darters were then still 
widespread in southern Ontario, apart from the 
Lake Ontario drainage, and possibly locally 
abundant in parts of many rivers (Figure 3a). D. E. 
McAllister and J. Kar (NMC 85-0564) collected one 
1.75 km northwest of Exeter, Ontario (43°24'45"N, 
81°29'50"W) in July 1985 in the Ausable River and 
sighted a few others. Much ofthis river is too heavily 
sedimented now for this species. 

Habitat 

Greatest abundance of Least Darters occurs in 
clear quiet, weedy waters of lakes and slow-moving 
rivers (Scott and Crossman 1973; Burr 1980; 
Kuehne and Barbour 1983). Muddy substrates or 
sand, or mixtures of the two, are found at these sites 
(Scott and Crossman 1973; Burr 1980; Page 1983). 
Trautman (1957) described preferred habitat as the 
clearest of waters with muck bottoms (= natural 
organic silt, not man-induced), debris, gravel, but 
not yellow clayey silts. 

These habitat types are probably widespread 
throughout the range of the species but could be 
threatened by development of water resources, such 
as drainage, channelization, construction of 



beaches, filling, soil tillage, and other alterations. 
Trautman (1957) assumed that such activities have 
caused a reduction in the abundance and range of 
the species in the United States prairies. 

Trends in the quality of habitat are unknown, but 
since the range of Least Darters in Canada is in one 
of the most heavily populated areas of the country, it 
is reasonable to suppose that the quantity of habitat 
may be decreasing. The disappearance of the species 
from Lake Ontario tributaries may be an example of 
such declines. 

General Biology 

The lifespan of Least Darters is quite short; as 
little as 18 months for males and 20 months for 
females (Burr and Page 1979). Both sexes are 
mature, and spawn, at 1 year of age (Burr and Page 
1979). Breeding in Canada probably occurs in May 
and June, although no published data on the subject 
has been found (Scott and Crossman 1973). 

Reproductive behaviour is unusual relative to 
other etheostomatins. Petravicz (1936) concluded 
from the numbers of each sex that he collected from 
shallow inshore waters and from deeper waters, that 
females stay in deep waters until ready to spawn 
while male darters enter shallow, heavily vegetated 
inshore waters during the breeding season. Petravicz 
(1936) noted that males do not establish territories, 
but do show resentment toward other males. Winn 
(1958b: as cited by Burr and Page 1979) observed 
that males defend small, three-dimensional 
territories, about 30 cm in diameter, although 
defense wasn't sustained through the night. 
Pugnacious behaviour between rival males, 
described by Burr and Page (1979), supports Winn's 
(1958b) observations. Winn (1958a) indicated that 
tank conditions affect the behaviour of many species 
of darters, and that crowding causes the breakdown 
of sex recognition in Etheostoma microperca. This 
variable behaviour may have caused these 
conflicting observations. 

In natural conditions, when a female is ready to 
spawn she will enter the shallow waters of the males' 
domain and make herself conspicuous (Petravicz 
1936). Several males attempt courting and a chase 
ensues while the female looks for a suitable site for 
egg deposition, usually a vertical or inverted surface 
on aquatic vegetation (Petravicz 1936; Burr and 
Page 1979). Eventually, one male, usually the largest 
and most colourful, succeeds in driving away his 
competitors (Petravicz 1936). When the female 
comes to rest motionless at her selected breeding 
site, the dominant male mounts her on the dorsal 
surface in a parallel position, using his enlarged 
pelvic fins as a clasping organ (Petravicz 1936; Burr 
and Page 1979). Vibration is initiated by the male 
but is quickly matched by the female (Petravicz 
1936). During the short lew seconds of vibration, the 



1990 



Dalton: Status of the Least Darter 



57 



bodies of each of the pair curve, a single adhesive egg 
is laid, and sperm are released near the egg 
(Petravicz 1936). Burr and Page (1979) found 
several eggs at each site, laid in three or four quickly 
repeated spawning acts. A few minutes after egg 
deposition, the female will find a new breeding site, 
and the pair will spawn again (Petravicz 1936). 

Spawning is continued from dawn till noon, or 
early afternoon, and each female lays about 30 eggs 
in one day (Petravicz 1936). Winn (1958b: as cited 
by Burr and Page 1979) reported pre-spawning 
female egg counts from 455 to 1102. No parental 
protection is provided to the eggs (Brooks and Page 
1979). 

Vertical and inverted spawning are remarkable 
feats for any fish, but especially for these darters 
which have no air bladder to control their buoyancy 
(Petravicz 1936). To maintain position on the 
breeding site, and on each other, Least Darter pairs 
rapidly vibrate their pectoral and caudal fins, and 
the male also vibrates his posterior dorsal fin 
(Petravicz 1936). Petravicz (1936) also observed 
horizontal spawning on the substrate but only in 
artificial conditions; aquaria that held few plants 
and a fine sand bottom. 

Food is chiefly small crustaceans and larval 
chironomids (Petravicz 1936; Burr and Page 1979; 
Paine et al. 1982). Other small benthic organisms 
found in stomachs of specimens examined by Burr 
and Page (1979) include gastropods, annelids and 
insects. The long caudal peduncle and pectoral fins, 
the laterally compressed body, and the small 
anteriorly opening terminal mouth are adaptations 
for capturing small, active prey, among plants 
(Paine et al. 1982). 

Burr and Page (1979) found no evidence of large- 
scale migration up or down stream. Winn (1958a) 
observed the populations of Least Darters in his 
study area, Whitmore Lake, Michigan, which is a 
lake with a muddy, organic debris laden bottom, 
moved from the deeper mucky areas to shallow 
shore water in April. This observation seems to 
imply that the species overwinters in deep, muddy 
water, when it is available, and agrees with 
Petravicz's (1936) observations on reproductive 
behaviour. 

Etheostoma microperca is hardier than many 
other fishes in drought conditions. The species 
survives extremely dry periods by burrowing into 
the substrate and waiting until more favourable 
conditions return (Tramer 1977: as cited by Burr 
and Page 1979). 



have been caused by competition between the two 
species (Pflieger 1971). Substrate type and current 
velocity exclude Least Darters from the Ozark 
lowlands (Pflieger 1971). Clear, upland streams are 
the natural habitat of the species. Large rivers, even 
those connecting the Ozarks and the northern range 
of the species, are barriers to dispersal of 
Etheostoma microperca (Pflieger 1971). 

Water temperature appears to limit the northern 
range of the species (Burr 1978). 

Habitat destruction, due to human activities, has 
caused population losses as well as reductions. 
Northern Ohio streams that have been dredged have 
suffered population losses or destruction (Trautman 
1957). Other human practices that have deleterious 
effects on fish habitats are listed by Pflieger (1971): 
ditching, drainage of swamps and other still waters, 
impoundments, channehzation, changing flow 
regimens. 

Pollutants from industrial and domestic 
activities, such as extensive pesticide and fertilizer 
use, also have deleterious effects on populations of 
Least Darter (Pflieger 1971). Shorehne development 
by cottagers, road construction etc., can cause 
increased soil erosion, resulting in higher turbidity 
levels, as well as reduction of the littoral zone of 
waterbodies, lessening the breeding area of Least 
Darters and the productivity of waterbodies in 
general. 

Special Significance of the Species 

Least Darters do not have any known economic 
value and their importance to the ecology of fresh- 
water systems is unknown and difficult to determine 
(Scott and Grossman 1973). Burr (1978) asserts that 
Etheostoma microperca is probably the most 
advanced member of the family Percidae and 
certainly the most advanced of the sub-genus 
Microperca. He recognized the species as 
monotypic but noted that several populations are in 
the process of genetic differentiation. These 
postulations may give the species importance in the 
study of evolutionary and speciation processes. 
Breeding habits are believed to warrant further 
study. Vertical and inverted spawning habits are 
unusual and may indicate a niche specialization 
(Scott and Grossman 1973). Winn (1958a) draws 
attention to the manner in which breeding habits 
change according to environmental conditions and 
states that the Least Darter's breeding behaviour is 
influenced in a complex manner by their size, as well 
as by environmental and genetic factors. 



Limiting Factors 

The distribution and range of Least Darters are 
controlled by several natural factors. In the Ozarks, 
the complementary distribution of Least Darters 
and Gypress Darters, Etheostoma proeliare, may 



Evaluation 

The range and numbers of Etheostoma micro- 
perca appear to have been reduced in parts of the 
United States. Trautman (1957) assumed that 
populations in Ohio were reduced because of 



58 



The Canadian Field-Naturalist 



Vol. 104 



habitat destruction. In 1972, Miller reported that the 
fish was rare in Kansas and endangered in 
Pennsylvania. More recently, Johnson (1987) stated 
that the species receives legal protection in Iowa and 
is of special concern in Kansas, Kentucky and 
Wisconsin. 

Least Darters are close to the northern fringe of 
their North American range in Canada. In the past 
several decades, the Canadian range appears to have 
contracted (Figure 3) due to habitat loss and 
degradation and, at present, the fish are found only 
in southwestern Ontario. Populations remaining in 
Canada appear to be healthy and stable and the 
range no longer seems to be shrinking. 

For the present, the species seems to be secure in 
Canada. However, its status should be re-evaluated 
in the near future, and, because of its small size and 
elusive habits, directed surveys are needed. 

Acknowledgments 

Many thanks are extended to G. Gale of OMNR 
and E. Holm of ROM for supplying information 
about collections of Least Darters by their 
respective organizations. I would especially Uke to 
thank D. E. McAllister, Canadian Museum of 
Nature, for providing collection record data as well 
as published literature concerning the species. R. R. 
Campbell is thanked for providing support and 
advice and R. E. Campbell for typing the 
manuscript. 

Literature Cited 

Burr, B. M. 1978. Systematics of the percid fishes of the 
subgenus Microperca, genus Etheostoma. Bulletin of 
the Alabama Museum of Natural History Number 4. 

Burr, B. M. 1980. Etheostoma microperca. In Atlas of 
North American freshwater fishes. Edited by D. S. Lee, 
C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. 
McAllister, and J. R. Stauffer, Jr. North Carolina 
Biological History Survey, Publication 1980-12. 

Burr, B. M., and L. M. Page. 1979. The life history of the 
Least Darter, Etheostoma microperca in the Iroquois 
River, Illinois. Illinois Natural History Survey 
Biological Notes Number 1 12. Urbana, Illinois. 

Holm, E., and E.J. Crossman. 1986a. Report on the 
search for an Ontario population of H. x-punctata and 



on a search for the species. Unpublished Royal Ontario 
Museum Report to Ontario Ministry of Natural 
Resources, Toronto, March 1986. 

Holm, E., and E. J. Crossman. 1986b. A report on the 
1985 attempt to resurvey some areas within the Ontario 
distribution of Clinostomus elongatus, the redside dace, 
and to summarize previous records. Unpublished Royal 
Ontario Museum report to Ontario Ministry of Natural 
Resources, Toronto. 

Johnson, J. E. 1987. Protected fishes of the United States 
and Canada. American Fisheries Society, Bethesda, 
Maryland. 

Kuehne, R. A., and R. W. Barbour. 1983. The American 
darters. The University Press of Kentucky, Lexington, 
Kentucky. 

Miller, R. R. 1972. Threatened freshwater fishes of the 
United States. Transactions of the American Fisheries 
Society 2: 239-252. 

Page, L. M. 1983. Handbook of darters. Illinois Natural 
History Survey, Champaign. THE Publications, 
Neptune City, New Jersey. 

Paine, M. D., J. J. Dodson, and G. Power. 1 982. Habitat 
and food resource partitioning among four species of 
darters (Percidae: Etheostoma) in a southern Ontario 
stream. Canadian Journal of Zoology 60: 1635-1641. 

Petravicz, J. J. 1936. The breeding habits of the Least 
Darter, Microperca punctulata Putnam. Copeia 
1936(2): 77-82. 

Pflieger, W. L. 1971. A distributional study of Missouri 
fishes. University of Kansas Publications, Lawrence, 
Illinois. 

Scott, W. B., and E. J. Crossman. 1973. Freshwater 
fishes of Canada. Fisheries Research Board of Canada 
Bulletin 184. Ottawa, Ontario. 

Smith, P. W. 1979. The fishes of Illinois. University of 
Illinois Press, Urbana, Illinois. 

Tramer, E.J. 1977. Catastrophic mortality of stream 
fishes trapped in shrinking pools. American Midland 
Naturalist 97(2): 469-478. 

Trautman, M. B. 1957. The fishes of Ohio. The Ohio 
State University Press, Columbus, Ohio. 683 pages. 

Winn, H. E. 1958a. Observations on the reproductive 
habits of darters (Pisces-Percidae). American Midland 
Naturalist 59(1): 190-212. 

Winn, H. E. 1958b. Comparative reproductive behav- 
iour and ecology of fourteen species of darters (Pisces- 
Percidae). Ecological Monographs 28(2): 155-191. 

Received 10 October 1989 



Status of the River Darter, Percina shumardi, in Canada* 



Ken W. Dalton 

8 Suffolk Street, Nepean, Ontario K2G 3P4 

Dalton, Ken W. 1990. Status of the River Darter, Percina shumardi, in Canada. Canadian Field-Naturalist 104(1): 
59-63. 

The River Darter, Percina shumardi, is a small percid of larger streams, rivers, and some inland lakes of Manitoba and 
northwestern Ontario, in Canada, as well as in the United States. The species is also known from the Lake St. Clair 
drainage of southwestern Ontario. Very little is known of the biology or ecology of the species in Canada and it does 
not seem to be abundant anywhere, although populations appear to be stable. 

Le Dard de riviere, Percina shumardi, est une petite espece de la famille des percides qui frequente les grands cours 
d'eau, les rivieres et certains lacs interieurs du Manitoba et du nord-ouest de I'Ontario au Canada, et I'espece est aussi 
trouve aux Etats-unis. On trouve egalement I'espece dans le bassin de drainage du lac Sainte-Claire dans le sud-ouest 
de I'Ontario. On connait tres peu de choses sur la biologic ou I'ecologie de I'espece au Canada et elle ne semble 
abondant nulle part, bien que les populations semblent stables. 

Key Words: River Darter, Percina shumardi, dard de riviere, Percidae, darters, rare and endangered fishes, status, 
Canada. 



River Darters, Percina shumardi {Girard 1860), 
are small fish (Figure 1) of the family Percidae 
(subfamily Etheostomatinae). They have slender 
but sturdy bodies and grow to about 58 mm total 
length in Canada (Scott and Grossman 1973). 
Longer lengths (to about 65 mm) have been 
reported for the fish in American waters (Cross 
1967; Thomas 1970; Smith 1979; Kuehne and 
Barbour 1983). The species has a large head with a 
short rounded snout and a moderately sized, 
terminal mouth (Trautman 1957; Scott and 
Crossman 1973). The large eyes are close together, 
high on the head (Kuehne and Barbour 1983). 

The dorsal fins are close together, the first 
having hard spines, while the second has soft rays 
(Trautman 1957; Scott and Crossman 1973), the 
caudal fin is slightly forked. Pelvic fins are of 
moderate size and are pointed and insert on the 
ventral side close behind the pectoral fins and are a 
small distance apart (Trautman 1957; Cross 1967). 
Each contains one "head" spine anteriorly, 
followed by five soft rays (Cross 1967). Pectoral 
fins are fan shaped and have soft rays but no 
spines. Two spines are found on the anal fin, 
followed by 11 to 13 rays (Scott and Crossman 
1973). 

The overall colouration of the River Darter 
varies from light brown to dark olive and many 
markings decorate the body. Seven or eight 
indistinct saddle marks adorn the dorsal surface 
and eight to ten dark lateral blotches appear on the 
sides, often as short vertical bars. Distinct 
suborbital bars drop from the eyes, and small, well- 



defined, caudal spots may be present (Scott and 
Crossman 1973). 

Colouration on breeding males is generally 
darker, and pigmented areas are more conspicu- 
ous. Posterior rays of the anal fin extend almost to 
the base of the caudal fin (Scott and Crossman 
1973; Kuehne and Barbour 1983). Breeding males 
may also develop nuptial tubercles on the caudal, 
anal and pelvic fins, as well as on the vent and on 
the head along the infraorbital and preopercular 
mandibular canals (Kuehne and Barbour 1983). 

Distribution 

River Darters have a very large latitudinal 
distribution (Figure 2), from Sipiwesk Lake in 
Manitoba, south to the Texas coast of the Gulf of 
Mexico (Scott and Grossman 1973: 1983). The 
Canadian range of the River Darter includes two 
provinces (Figure 3); collection records of the 
National Museum of Natural Sciences, now 
Canadian Museum of Nature (NMG), the Ontario 
Ministry of Natural Resources (OMNR), the 
Royal Ontario Museum (ROM) and the Province 
of Manitoba were accessed, in addition to 
literature records. 

In Ontario, it occurs from the Kenora district to 
as far north as Lake Attawapiskat and Sandy Lake 
(Scott and Crossman 1973). The species is also 
known from the Lake St. Clair drainage: a 
Sydenham River collection was made in 1975 
(OMNR 582) and one from St. Luke's Bay in Lake 
St. Glair itself (ROM 48870) in 1985. 

In Manitoba, the species occurs in scattered 
locations in the Lake Winnipeg drainage as far 



♦Reviewed and accepted by COSEWIC 1 1 April 1989 — no designation required. 

59 



60 



The Canadian Field-Naturalist 



Vol. 104 




lOmm 
Figure L River Darter, Percina shumardi {adapted from Cross 1967). 



north as Sipiwesk Lake and as far west as Lake 
Dauphin and the Red River (Scott and Grossman 
1973). 

Protection 

River Darters are legally protected in the State 
of Ohio but not elsewhere in the United States, 
although the species is given special concern in 
Kansas, Kentucky, North Dakota, and West 
Virginia (Johnson 1987). In Canada, Percina 
shumardi have no specific protection, but 
incidental protection is provided by the Fish 
Habitat Section of the Fisheries Act. 

Population Sizes and Trends 

Percina shumardi does not seem to have ever 
been abundant in Canadian waters (Scott and 
Crossman 1973). Not many collections have been 
made in Canada (NMC, OMNR, ROM records), 
and usually only one or two fish have been taken 
from a site, the most in a single collection being 10 
specimens. The fish's range includes some 
relatively remote areas of the country and the 
paucity of collection records may be a reflection of 
this. It may also be a result of difficulty 
encountered in sampling the species' favoured 
habitat, large rivers with fair to moderate currents 
and a rocky substrate. 

Most Ontario records of River Darters are from 
lakes in the northwest of the Province; only two 
Ontario records actually come from rivers 
(Vermilion Creek, ROM 07709; Sydenham River 
(east), OMNR 582). The latter, and the collection 
from St. Luke's Bay, may indicate a range 
expansion into Lake St. Clair and southern Ontario. 
Scott and Crossman ( 1 973) had earlier suggested the 
possibility of a southwestern Ontario distribution 
based on records from the Detroit River. 

Actual numbers of River Darters in Canadian 
waters cannot be estimated. The species seems to be 
naturally rare in Ontario and Manitoba, although 



they may exist undetected in many rivers. Records, 
although few and far between, seem to indicate 
stable populations in both provinces. 

In the United States, the distribution is 
widespread in the Mississippi system but the species 
has never been detected in abundance (Trautman 
1957). The status of the species is considered to be of 
special concern in most U.S. states where it occurs 
(Johnson 1987). 

Habitat 

River Darters are chiefly found in large rivers 
with rocky substrates and fair to moderate currents 
(Thomas 1970; Scott and Crossman 1973). Large 
individuals have also been taken from strong 
currents (Page 1983). Thomas (1970) collected the 
fish mostly from deep riffles and chutes, and only 
found young fish in water 60 cm deep or shallower. 
River Darters also inhabit lakes (Scott and 
Crossman 1973) and in Ontario are more commonly 
found in lacustrine environments. Smith (1979) 




FiCiURK 2. North American range of the River Darter, 
Percina shumardi (adapted from Kuehne and 
Barbour 198.1). 



1990 



Dalton: Status of the River Darter 



61 



100 100 200 300 400 

SCALE IN MILES 




B 



Figure 3. Distribution of the River Darter, Percina shumardi, in Canada; A - collections 
pre-1975, B - collections post-1975. 
• Ontario Ministry of Natural Resources (OMNR) 
■ Royal Ontario Museum (ROM) 
A Canadian Museum of Nature (NMC) 
n Province of Manitoba 



62 



The Canadian Field-Naturalist 



Vol. 104 



reports the fish infrequently occurs in clear, sandy 
channels close to large rivers. 

General Biology 

No information about the spawning behaviour 
of River Darters in Canada is available. On the 
Neoshi River, Kansas, Cross (1967) captured River 
Darters that seemed to be spawning below a low 
dam in clear water about 61 cm deep, over a 
bedrock substrate littered with gravel and large 
stones. Scott and Crossman (1973) presumed that, 
in Canada, breeding occurs in the the spring, 
possibly during the months of June and July. The 
authors also assumed that spawning behaviour is 
similar to that of other members of the genus 
Percina, P. copelandis and P. maculata in 
particular. Breeding of these two species, as 
reported by Scott and Crossman (1973), has 
several common features that may be shared by 
River Darters. The female wriggles her body into 
gravel and then the male perches on top of her, 
resting his caudal fin beside hers. Eggs are laid, 
fertilized, and deposited from that position and 
given no parental care. The fish are promiscuous 
and females will mate with several partners during 
the breeding season. 

The breeding age of the species, and the number 
of eggs laid by River Darters, are unknown. 

No direct observations of feeding by River 
Darters have occurred in Canada. Thomas (1970) 
found that, in the lower Kaskasia River, Illinois, 
the fish principally ate chironomid and hydropsy- 
chid larvae and also took other insects, microcrus- 
taceans, and fish eggs. Starmes (1977, as cited by 
Kuehneand Barbour 1983) and Thomson (1974, 05 
cited by Page 1983) state that snails, when 
available, from a large part of the diet of P. 
shumardi. 

The age attained by River Darters in Canada is 
unknown; most of those in Illinois die at three 
years of age, although some may reach four years 
(Smith 1979). 

Movement by River Darters is generally 
unknown; however, twice, when the Scioto River, 
Ohio, was in flood, Trautman (1956) captured a 
single adult in flood water a good distance from the 
river channel, and suggested that they were 
migrating upstream. 

Limiting Factors 

The River Darter is believed to be limited to 
large streams and lakes (Thomas 1970; Kuehne 
and Barbour 1983). It is more tolerant of turbidity 
than other darters (Pfleiger 1975), and even seems 
to prefer shallow water in turbid locations 
(Trautman 1957). 



Pollution does not appear to be a major limiting 
factor in the western Canadian range of River 
Darters, but could be limiting in southwestern 
Ontario. Thomas (1970) reports a few dramatic 
cases where pollution killed many fish, including 
River Darters, in Illinois. Impoundments have led 
to the extirpation of the fish from some American 
rivers (Thomas 1970; Kuehne and Barbour 1983) 
as have dredging and channelization (Thomas 
1970). 

Special Significance of the Species 

River Darters span a large climatic and 
latitudinal range, from the Gulf of Mexico in 
Texas to Sipiwesk Lake in northern Manitoba 
(Kuehne and Barbour 1983). They have no known 
public interest or economic importance. 

Many darters of the genus Percina are of special 
concern, and some are legally protected in various 
U.S. states {see Johnson 1987). 

Evaluation 

River Darters appear to be rare in Canada, and 
although populations seem to be stable, their 
status could change rapidly if habitat destruction 
occurs. Populations should be monitored to detect 
change, but at present the species does not appear 
to be under any immediate threat nor to warrant 
vulnerable status. 

Acknowledgments 

Thanks are due to G. Gale of OMNR and E. 
Holm of ROM for providing collection record 
data and for help in verifying questionable records; 
to D. E. McAllister of the Canadian Museum of 
Nature for providing collection records and copies 
of much of the literature cited in this report. R. R. 
Campbell, Department of Fisheries and Oceans, 
also deserves thanks for encouragement and advice 
during the writing of this report. 

Literature Cited 

Cross, F. B. 1967. Handbook of fishes of Kansas. 
Museum of Natural History, University of Kansas, 
Lawrence, Kansas. 

Johnson, J. E. 1987. Protected fishes of the United 
States and Canada. American Fisheries Society, 
Bethesda, Maryland. 

Kuehne, R. A., and W. A. Barbour. 1983. The Ameri- 
can darters. University Press of Kentucky, Lexington, 
Kentucky. 

Page, L. M. 1983. Handbook of darters. THF Publica- 
tions, Neptune City, New Jersey. 

Pfleiger, W. L. 1971. A distributional study of Missouri 
fishes. Mu.seum of Natural History, University of 
Kansas, Lawrence, Kansas. 



1990 



Dalton: Status of the River Darter 



63 



Scott, W. B., and E.J. Grossman. 1973. Freshwater 

fishes of Canada. Fisheries Research Board of Canada 

Bulletin 184. 
Smith, P. W. 1979. The fishes of Illinois. University of 

Illinois Press. Urbana, lUinois. 
Starmes, W. C. 1977. Theecology and life history of the 

endangered snail darter, Percina (Imostoma) tanasi. 

Knoxville: University of Tennessee Wildlife Resources 

Agency Technical Report 77-52. 
Thomas, D. L. 1970. An ecological study of four 

darters of the genus Percina (Percidae) in the 



Kaskaskia River, Illinois. Illinois Natural History 
Survey Biological Notes, No. 70. 18 pages. 

Thompson, B. A. 1974. An analysis of sympatric 
populations of two closely related species of Percina, 
with notes on food habits of the subgenus Imostoma. 
ASB Bulletin 21: 87. 

Trautman, M. B. 1957. The fishes of Ohio. Ohio State 
University Press. Baltimore, Maryland. 



Accepted 10 October 1989 



Status of the Redbreast Sunfish, Lepomis auritus, in Canada* 

J. Houston 

374 Fireside Drive, Woodlawn, Ontario KOA 3M0 

Houston, J. 1990. Status of the Redbreast Sunfish, Lepomis auritus, in Canada. Canadian Field-Naturahst 104(1): 
64-68. 

The Redbreast Sunfish, Lepomis auritus, is one of the smaller centrarchids. The native range is confined to the Atlantic 
drainage east of the Appalachians. In Canada, the species is found only in New Brunswick where it has a restricted 
distribution. It is often overlooked because of its small size and its similarity to smaller members of other centrarchid 
species. Due to the restricted range and the potential for habitat pollution by industrial, urban, agricultural and 
silvicultural contaminants, the species should be considered vulnerable in Canada. 

Le crapet rouge, Lepomis auritus, est I'un des centrarchides les plus petits. Le territoire de I'espece est limite au bassin 
versant de I'Atlantique a Test des Appalaches. Au Canada, on ne trouve le crapet rouge qu'au Nouveau-Brunswick ou 
son aire de repartition est restreinte. Ce poisson passe souvent inaper9u en raison de sa petite taille et de sa 
ressemblance avec les plus petits membres d'autres especes de centrarchides. A cause de son territoire limite et des 
risques de contamination des bassins versants qu'il occupe par des poUuants industriels, urbains, agricoles et forestiers, 
I'espece doit etre consideree comme vulnerable au Canada. 

Key Words: Redbreast Sunfish, Lepomis auritus, crapet rouge, Centrarchidae, sunfishes. New Brunswick, rare and 
endangered fishes, status, Canada. 



The Redbreast Sunfish, Lepomis auritus 
(Linnaeus 1758), is one of the smaller members of 
the sunfish family (Centrarchidae). These fish 
(Figure 1) usually range from 12.7 to 17.8 cm in 
length but individuals as large as 20.3 cm have 
been reported from New Brunswick and 23.9 cm in 
Connecticut (Scott and Crossman 1973). The large 
Connecticut individual weighed 312 g but the 
usual range is around 165 to 232 g (see Carlander 
1977). Redbreast Sunfish have a deep, laterally 
compressed body with a narrow caudal peduncle. 
The distinctive opercular flap is larger in adult 
males than in juveniles or adult females and is 
black with no distinctive border. 

The body is golden brown to olivaceous, the 
dorsal surface being darker. The sides are lighter in 
colour showing indistinct red spots and blue 
streaks, particularly on the side of the head. The 
breast, as the name implies is usually red, but may 
vary from yellow to orangish-red. The colour is 
intensified in breeding males. Fins may be dusky or 
darkly mottled, the leading edges and tips of the 
pectorals are dark {see Scott and Crossman 1973). 

The species may be confused with the Longear 
Sunfish (Lepomis megalotis) or the Pumpkinseed 
(Lepomis gibhosus), but the longer opercular flaps 
of adult male Redbreast Sunfish are not bordered 
with white or colour. Females and smaller 
individuals may also be identified by the presence 
of palatine teeth which are not found in the 



Longear Sunfish (Scott and Crossman 1973). 
However, the ranges of the Redbreast and Longear 
sunfishes do not overlap in Canada. 

As an economically unimportant sunfish the 
species is often overlooked, especially where larger 
species occur in the range. This, along with 
confusion with the Longear Sunfish may lead to 
erroneous assumptions regarding its abundance. 

Distribution 

The Redbreast Sunfish is one of the few native 
lepomin sunfishes occurring east of the Appalach- 
ians (Lee 1980). The native range (Figure 2) 
extends from New Brunswick (in Canada) south 
along the Atlantic slope to central Florida. The 
species is not normally found west of the 
Appalachians and in the south the native range 
extends west of the Choctawhatchee and 
Apalachicola rivers (Scott and Crossman 1973; 
Lee 1980). The Redbreast Sunfish has not been 
recorded from the state of Mississippi, but has 
been introduced into several lakes outside of the 
normal range, particularly in Texas and Oklahoma 
(Scott and Crossman 1973) and also northern Italy 
(Lee 1980). 

In Canada, the species is known only from New 
Brunswick where it has been found in the Canaan, 
Oromocto, Magaguadavic, and Kennebecasis 
river systems as well as Anne, Oromocto, and 
Yoho lakes (Gorham 1970; Scott and Crossman 



♦Vulnerable status approved and assigned by COSEWiC 1 I April 1989. 



64 



1990 



HOUSTON: Status of the Redbreast Sunfish 



65 




Figure 1. Redbreast Sunfish, Lepomis auritus [drawing by P. Buerschaper, from Scott 
and Grossman (1973) by permission]. 



1973). It may have a wider distribution, but 
because it is small and may readily be confused 
with other sunfishes, particularly the Pumpkin- 
seed, its correct identification could have often 
been overlooked. One would expect, however, that 
serious collectors would have noted its presence. 

Protection 

No specific protection measures are offered the 
species although general protection is given under 
the Fisheries Act and the New Brunswick 
Provincial Endangered Species Act. 

Population Sizes and Trends 

Very little information is available on the 
distribution and abundance of the Redbreast 
Sunfish in New Brunswick. Collection records for 
the species are presented in Table 1. It has been 
suggested that the species has been known in New 
Brunswick since at least 1896 (Cox 1896). The first 
authentic record dates from 1948 (Scott and 
Crossman 1973). 

The rarity of records may be indicative of 
natural rarity of a peripheral species, or could be 
due to lack of interest in the species and confusion 
with the Pumpkinseed. The more probable 
situation is likely to be a combination of these 
factors. No recent records (1980 on) are indicated 
but the species has definitely been taken in 
Oromocto Lake at various times between 1960 and 
1979. 

Habitat 

The Redbreast Sunfish appears to be adaptive to 
a wide range of habitats. In New Brunswick, the 
species inhabits streams and lakes with rocky 
bottoms. In streams they occupy the slower deeper 
areas over rocks and gravel, in lakes they occur 



most abundantly on rocky shoals or in deeper, 
quieter water with sand or mud bottoms and 
emergent vegetation (Scott and Crossman 1973). 
The species tends to be more of a riverine species 
than other sunfishes (Lee 1980) and the description 
of nesting sites in Pennsylvania (Buynak and Mohr 
1978) indicates that habitat requirements there are 
similar to those in New Brunswick. However, in 
Georgia the species frequents streams of the 
coastal plain characterized by low pH (4.5 to 6.0) 
with periods of extreme high and low flows 
(Coomer et al. 1977). Bottom substrates are 
usually of sand, sandstone and rubble. 

In summer the fish appear to be widely 
distributed but aggregate in schools in deeper 
waters for the winter when water temperature falls 
below 5°C (Breder and NigreUi 1935; Scott and 
Crossman 1973). In spring they migrate to shallow 
areas of lakes (15 to 46 cm) or downstream of 
rapids in streams for spawning when water 
temperatures go above 16.7° C, dispersing for the 
summer following spawning. The species has been 
reported to spawn in brackish, tidal waters in some 
locations where water temperature, salinity and 
depth vary considerably (Richmond 1940). 

General Biology 

There is very little published on the biology of 
the species in Canada. Scott and Crossman (1973) 
have summarized information on the biology and 
ecology of the species in the northern parts of the 
range. Redbreast Sunfish spawn in the spring or 
early summer when water temperatures are above 
1 6° C (Scott and Crossman 1973). They overwinter 
in schools (Breder and Nigrelli 1935) in areas of 
deeper water and begin to move to shallower 
waters when water temperature rises above 10°C 
(Breder 1936; Breder and Rosen 1966). Spring 



66 



The Canadian Field-Naturalist 



Vol. 104 




Figure 2. Native North American range 
Redbreast Sunfish {Lepomis auritus). 



spawning migrations appear to be differentiated by 
sexes, the males making their way to the shallows 
first. The males establish territories and build nests 
(redds), which may be 60 to 100 cm in diameter and 
in water 15 to 46 cm in depth (Scott and Grossman 
1973). Females leave the area after the eggs are 
laid, the males remain to guard the redd, fanning 
the nest to prevent suffocation and possibly 
guarding the larvae until swimup (Scott and 
Grossman 1973). 

In lacustrine areas the redds may be close 
together in the open, but in streams they are in the 
current downstream of a protecting rock. They 
may also utilize unused redds of other centrarchids 
(Scott and Grossman 1973). Not surprisingly, 
Redbreast Sunfish have been known to hybridize 
with other centrarchids {see Garlander 1977) 
including the Bluegill {Lepomis macrochirus), the 
Redear Sunfish {Lepomis microlophus). Green 
Sunfish {Lepomis cyanellus), and the 
Pumpkinseed. 

The number of eggs per female increases with 
age and size, but ranges from 1000 to 8000 {see 
Garlander 1977). The eggs are about 2 mm in 
diameter, amber in colour, and are adhesive (Scott 
and Grossman 1973; Garlander 1977; Buynak and 
Mohr 1978). Fertilized eggs collected from the wild 
hatched in three days when retained at 20 to 24° G 
(Buynak and Mohr 1978). Newly hatched larvae 
averaged 4.9 mm in length and by swimup 
averaged 7.9 mm. Other information and descrip- 
tions of Redbreast Sunfish larvae are also 



mentioned by Buynak and Mohr (1978). Age, 
weight and length data are presented by Garlander 
(1977). Growth appears to be rapid, not sexually 
differentiated, and maturity is reached at about 

23 g in the second year {see Garlander 1977). 
Growth is apparently slower in the northern parts 
of the range (Garlander 1977). In New Brunswick, 
maximum size is about 20 cm, but the average is 13 
to 18 cm with approximate weights of 165 to 232 g. 
In Gonnecticut, they are said to be larger, up to 

24 cm in length and 312 g in weight (Scott and 
Grossman 1973; Garlander 1977). Longevity is not 
known, but individuals to age eight have been 
recorded (Garlander 1977). 

Redbreast Sunfish feed primarily on small 
aquatic insects, but adult insects, molluscs, benthic 
invertebrates and even small fish may be consumed 
(Scott and Grossman 1973; Goomer et al. 1977). 
The species actively selects chironomids through- 
out the year, but are opportunistic feeders taking 
anything of the proper size which is available 
(Goomer et al. 1977). Lauder (1980) has described 
the feeding mechanism of these and other 
sunfishes. 

Brown Bullhead {Ictalurus nebulosus), Ghain 
Pickerel {Esox niger) and American Eel {Anquilla 
rostratra) are the main predators in the Ganadian 
range (Scott and Grossman 1973). In other parts of 
the range the species may be prey to any of the 
larger piscivores present. Many fishes will take the 
eggs from unguarded nests (Scott and Grossman 
1973). Hoffman (1967) has listed parasitic 
infections. 

Limiting Factors 

There is no information available on limiting 
factors for this species which appears adaptable to 
a wide range of ecological conditions. They do well 
in water of pH 4.8 to 8.4, salinity to 8% and water 
temperature to 37° G {see Garlander 1977). They 
tend to do better in uncrowded conditions with few 
predators. 

The species requires clean water and, like most 
other fishes, does not fare well in polluted waters. 
No reliable information of the effects of aquatic 
contaminants on the species in the New Brunswick 
range is available, but these waters are subject to 
chemical pollution from industry (particularly the 
pulp and paper industry), agricultural runoff, 
urban wastes and pesticides (silviculture insect 
control programs). 

Special Signiflcance of the Species 

Due to its small size the species is one of the least 
important (to man) sunfishes. In Ganada and New 
Brunswick, lepomin sunfishes and other warm 
water fishes are not highly regarded as food or 
sport fish. However, they will take live bait and if 



1990 



HOUSTON: Status of the Redbreast Sunfish 



67 



Table 1. Collection records of Lepomis auritus in Canada (New Brunswick) to March 1988. ROM = Royal Ontario 
Museum, NBM = New Brunswick Museum, NMC = National Museum of Natural Sciences, NBNRDS = New 
Brunswick Department of Natural Resources Data System. 













Number in 


Catalogue # 


Location 


Latitude 


Longitude 


Date 


Collection 


— 


Canaan River 


— 


— 


03/09/48 


— 


ROM 22087 


Oromocto Lake 


45°36'90"N 


67°00'90"W 


14/08/61 


— 


ROM 22085 


Oromocto River 


— 


— 


24/06/62 


— 


NBM 0001 


Oromocto River 


45°36'00"N 


65°55'00"W 


1960 


1 


NBM 0011 


Oromocto River 


45°5r00"N 


66°29'00"W 


1960 


1 


NBM 0119 


Kings County 


— 


— 


30/05/66 


— 


NBM 0399 


Rusagonis Stream 


45°49'00"N 


66°32'00"W 


30/07/67 


10 


NBM 0494 


Rusagonis 


45°48'00"N 


61°37'00"W 


25/05/68 


4 


NBM 1068 


Oromocto Lake 


45°24'00"N 


66°38'00"W 


1979 


— 


NMC 67-0015 


Anne Lake 


45°25'00"N 


66° 13 '00" W 


15/07/66 


4* 


NMC 67-0322 


Anne Lake 


45°25'00"N 


66°13'00"W 


28/08/67 


11 


NMC 68-0072 


Anne Lake 


45°25'00"N 


66°13'00"W 


03/07/67 


3 


NBNRDS 


Oromocto Lake (St. John River) 






P** 


NBNRDS 


Yoho Lake (St. John River) 






P 


NBNRDS 


Modsley Lake (St. Croix River) 






P 


NBNRDS 


Knockdrin Lake (East Musquash River) 






P 


NBNRDS 


Shadow Lake (East 


Musquash River) 






P 



*Also catalogued by NBM as 0155. 
'=*P = Present. 



fished with hght tackle, can provide interesting 
fishing (Scott and Grossman 1973). The fish are 
good test animals and can be easily maintained in 
captivity (Carlander 1977). It is doubtful that they 
would ever become a popular aquarium fish. Some 
may be taken incidentally from the lower Saint 
John River and sold in the Farmer's Market at 
Saint John (M.A. Redmond, New Brunswick 
Department of Natural Resources, Fredericton, 
New Brunswick; personal communication). 

Evaluation 

The Redbreast Sunfish appears to inhabit the 
northern fringe of its North American range in 
New Brunswick. The species is restricted to 
drainages of the Atlantic seaboard but has been 
successfully introduced into areas outside of the 
native range. Although populations are locally 
abundant (Scott and Grossman 1973), they are 
known in Canada from only a few localities in New 
Brunswick where there is a high potential of 
environmental risk from water borne contami- 
nants resulting from industrial, agricultural and 
silvicultural use of toxic chemicals. Due to the very 
restricted range and the potential risk of 
contamination in the watersheds where the species 
exists, the Redbreast Sunfish should be considered 
'Vulnerable' in Ganada. 

Acknowledgments 

This report was made possible through the 
support of the Department of Fisheries and 



Oceans, the Canadian Wildlife Service, and World 
Wildlife Fund Ganada. The cooperation of Don 
McAllister of the National Museum of Natural 
Sciences (now Canadian Museum of Nature) in 
preparation and review of the report and provision 
of collection records is gratefully acknowledged. 
The staff of the Royal Ontario Museum, the New 
Brunswick Museum, and the New Brunswick 
Department of Natural Resources are also 
acknowledged for their assistance in the provision 
of collection records and helpful advice. 

Literature Cited 

Breder, C. M., Jr. 1936. The reproductive habits of the 
North American sunfishes (family Centrarchidae). 
Zoologica 21(1): 1-48 + plate. 

Breder, C. M., Jr., and R. J. Nigrelli. 1935. The influence 
of temperature and other factors on the winter 
aggregation of the sunfish, Lepomis auritus, with critical 
remarks on the social behaviour of fishes. Ecology 16(1): 
33^7. 

Breder, C. M. Jr, and D. E. Rosen. 1966. Modes of 
reproduction in fishes. Natural History Press, New 
York, New York. 

Buynak, G. L., and H. W. Mohr, Jr. 1978. Larval 
development of the Redbreast Sunfish {Lepomis 
auritus) from the Susquehanna River. Transactions of 
the American Fisheries Society 107(4): 600-604. 

Carlander, K. F. 1977. Handbook of freshwater fishery 
biology. Volume 2. Iowa State University Press, Ames, 
Iowa. 

Coomer, C. E., Jr., D. R. Holder, and C. D. Swanson. 
1977. A comparison of diets of Redbreast Sunfish and 
Spotted Suckers in a coastal stream. Proceedings of the 
Annual Conference of the Southeastern Association of 
Fish and Wildlife Agencies 31: 587-596. 



68 



The Canadian Field-Naturalist 



Vol. 104 



Cox, P. 1896. Catalogue of the marine and freshwater 

fishes of New Brunswick. Bulletin of the Natural History 

Society of New Brunswick 13: 62-75. 
Gorham, S. W. 1970. Distributional checklist of the 

fishes of New Brunswick. New Brunswick Provincial 

Museum, St. John, New Brunswick. 
Hoffman, G. L. 1967. Parasites of North American 

freshwater fishes. University of California Press, Los 

Angeles, California. 
Lauder, G. V. 1980. The suction feeding mechanism in 

sunfish (Lepomis): an experimental analysis. Journal of 

Experimental Biology 88: 49-72. 
Lee, D. S. 1980. Lepomis auritus. Redbreast Sunfish. 

Page 590 in Atlas of North American freshwater fishes. 



Edited by D. S. Lee, C. R. Gilbert, C. H. Hocutt, R. E. 

Jenkins, D. E. McAUister, and J. R. Stauffer, Jr.. North 

Carolina State Museum of Natural History, Raleigh, 

North Carolina. Biological Survey Publication Number 

1980-12. 
Richmond, N. D. 1940. Nesting of the sunfish, Lepomw 

auritus (Linnaeus), in tidal water. Zoologica 23(3): 

329-331. 
Scott, W. B., and E.J. Grossman. 1973. Freshwater 

fishes of Canada. Fisheries Research Board of Canada 

Bulletin 184. 



Accepted 10 October 1989 



Status of the Orangespotted Sunfish, Lepomis humilis, in Canada* 
Douglas B. Noltie 

School of Natural Resources, Fisheries and Wildlife Program, 112 Stephens Hall, University of Missouri — 
Columbia, Columbia, Missouri 65211. 

Noltie, Douglas B. 1990. Status of the Orangespotted Sunfish, Lepomis humilis, in Canada. Canadian 
Field-Naturalist 104(1): 69-86. 

The Orangespotted Sunfish, Lepomis humilis, has only recently been described as a member of the fish fauna of Canada. 
Given this novelty, a detailed description of the species is provided. Some morphological distinctions exist between 
specimens from Canadian and American populations. Although common and widely distributed in the United States, its 
known range in Canada is restricted to only three locations in southwestern Ontario. Evidence presented suggests that the 
species is native to the Lake Erie basin but that Canadian populations may have arisen through direct range expansion 
northwards out of Ohio. Population estimates and trends have not been established, but prospects for the species' long- 
term establishment in Ontario appear good given trends in southern populations. The species' habitat in Ontario is 
primarily low-gradient, slow-flowing streams having turbid waters, silt, clay or mud bottoms, and minimal submerged 
aquatic vegetation, a feature shared with more southerly populations. The changing face of southwestern Ontario stream 
habitats has favoured their invasion, and continuing agricultural impact bodes well for the numerical and geographical 
increase of the species in Canada. At present, the species is subject to no specific protection. A species of great beauty 
among Canadian fishes, a designation of vulnerable is suggested given its low numbers and extremely limited distribution. 

Le crapet menu, Lepomis humilis, est presente comme une espece relativement nouvelle parmi les poissons du Canada. 
Etant donne qu'il s'agit d'une nouveaute, on donne une description detaillee de I'espece. Certaines differences 
morphologiques existent entre les specimens provenant des populations canadiennes et americaines. Bien que ce poisson 
soit largement repandu aux Etats-Unis, son aire de dispersion connue au Canada est limitee a trois endroits seulement au 
sud-ouest de I'Ontario. Les faits presentes portent a croire que I'espece est indigere au bassin du lac Erie mais que ces 
populations canadiennes semblent etre apparues grace a une expansion directe de I'aire de dispersion vers le nord a partir 
de I'Ohio. On n'a pas etabli d'estimations ni de tendance des populations, mais les perspectives d'etablissement a long 
terme de I'espece en Ontario semblent bonnes, etant donne les tendances observees dans les populations du sud. En 
Ontario, cette espece frequente surtout des cours d'eau turbides a courant faible, et a inclinaison peu prononcee dont le 
fond est constitue de limon, d'argile ou de vase et ou il y a un minimum de vegetation aquatique submergee, 
caracteristiques qu'elle partage avec les populations plus meridionales. Les changements de I'habitat des cours d'eau du 
sud-ouest de I'Ontario ont favorise leur invasion et les repercussions constantes de I'agriculture augurent bien pour 
I'expansion numerique et geographique de I'espece au Canada. Pour le moment, I'espece ne beneficie d'aucune protection 
particuliere. On propose de classer cette espece, d'une grande beaute parmi les poissons du Canada, comme vulnerable 
donne etant le nombre peu eleve d'individus et leur distribution exremement limitee. 

Key Words; Orangespotted Sunfish, Lepomis humilis, crapet menu, Centrarchidae, sunfishes, Ontario, rare and 
endangered fishes, distribution, population size and trends, range expansion, ecology, habitat, reproduction, life 
history, status, Canada. 

The Orangespotted Sunfish, Lepomis humilis Description 
(Girard 1858), has only recently been described in The Orangespotted Sunfish is one of the most 

Canada (Holm and Coker 1981). Some evidence briUiantly coloured of the Lepomis species (Eddy 

(Noltie and Beletz 1984) suggests that these and Underbill 1974). Specimens (Figure 1) are 

populations have arisen in southwestern Ontario moderately deep and compressed to somewhat 

through a direct range extension northwards out of oblong in shape (Bollman 1892; Hay 1894; Jordan 

Ohio. Since the present Canadian distribution is and Evermann 1902; Forbes and Richardson 1920; 

extremely restricted, and populations exist in low Cross 1967; Becker 1983) and of rather slender build 

numbers in an area subject to intensive agriculture, (Eddy and Underbill 1974). Each possesses 

this report was commissioned to bring the species to relatively high dorsal spines and a long, broad 

the attention of COSEWIC. Given the species' opercular flap (Jordan and Evermann 1902) which 

novelty and the fact that some morphological is flexible (Richardson 1904; Eddy and Surber 1947; 

distinctions exist between populations in the United Holm and Coker 1981). The conspicuous black 

States and Canada (Holm and Coker 1981), a patch this flap bears has a coloured margin varying 

detailed description is provided. from white (Harlan and Speaker 1956; Moore 1957; 



♦Vulnerable status approved and assigned by COSEWIC 1 1 April H 

69 



70 



The Canadian Field-Naturalist 



Vol. 104 




Figure 1. Orangespotted Sunfish, Lepomis humilis, from Canard River, Ontario, June 
1983 (from Noltie and Beletz 1984). 



Cross 1967; Miller and Robison 1973; Buss 1974; 
Pflieger 1975; McClane 1978; Becker 1983) to 
greyish white (Barney and Anson 1923), white with 
an occasional flush of orange (Trautman 1981), pale 
with tinges of red or orange (Eddy and Surber 1947; 
Eddy and Underbill 1974), wide and pale lavender, 
pinkish or light crimson (Richardson 1904; Forbes 
and Richardson 1920), to broad and red, pinkish or 
orange (Thompson and Hunt 1930; Eddy 1957; Clay 
1962) or broad and red, entirely surrounding the 
black (Hay 1894; Jordan and Evermann 1896, 
1904). The mouth is large, the posterior of the 
maxillary extending beyond the anterior orbit rim 
(Girard 1858; Moore 1957; Cross and ColUns 1975). 
The scales of the throat are much smaller than those 
on the flanks (Clay 1962). The morphology of their 
pharyngeal bones and teeth has been outlined by 
Richardson (1904). 

Typical body colour is olive to bluish, with gold to 
green spots or motthngs posteriorly (Hay 1894; 
Jordan and Evermann 1896, 1902; Richardson 
1904; Forbes and Richardson 1920; Harlan and 
Speaker 1956; Cross 1967). The sides and back are 
greenish with silver-blue reflections, and the belly is 
white or yellow (Pflieger 1975; Becker 1983), or 
orange to red as are the lower fins (Bollman 1 892; 
Hay 1894; Jordan and Evermann 1896, 1902; 
Forbes and Richardson 1920; Eddy and Underbill 
1974). The anal fin has a dusky distal margin 
(Forbes and Richardson 1920). Often a faint black 
dot occurs on the last rays of the dorsal fin (Jordan 
and Evermann 1896). The cheeks and opercules 
possess longitudinal radiating brown, rusty orange, 
orange, golden, red or emerald streaks (Forbes and 
Richardson 1920; Barney and Anson 1923; 
Thompson and Hunt 1930; Eddy and Surber 1947; 
Clay 1962; Eddy and Underbill 1974; McClane 
1978). Bright red, salmon-red, or orange spots 
numbering 20 to 30 are scattered irregularly over the 
sides of males, but those on females are brown (Hay 
1 894; Jordan and Evermann 1896, 1902; Forbes and 



Richardson 1920; Harlan and Speaker 1956; Clay 
1962; Buss 1974; Eddy and Underbill 1974; 
McClane 1978; Holm and Coker 1981). As such, the 
Orangespotted Sunfish is one of the few sexually 
dichromatic centrarchids (Becker 1983; Petersen et 
al. 1986), colour differences alone being used to sex 
individuals with 95% accuracy (Petersen 1979; 
Petersen et al. 1986). 

In colouration, non-breeding males are generally 
silvery with darker backs and whitish bellies (Miller 
1963; Trautman 1981). During breeding, the males 
become more vividly coloured, taking on a pattern 
more complicated than in most sunfishes (Breder 
1936), and thereby differentiating the sexes (Breder 
and Rosen 1966). A breeding male is opalescent 
green with the sides of the head iridescent pale-blue 
streaked by bright orange, the breast and belly 
orange or red, the fins brilliantly orange, the dorsal 
fin rays tipped with crimson, the pelvics and anal 
outlined by black, and the irises red or orange 
(Barney and Anson 1923; Cook 1959; Cross 1967; 
Miller and Robison 1973; Trautman 1981; Becker 
1983). The belly and lower fins deepen in colour also 
(Harlan and Speaker 1956). The dorsal in males may 
possess a narrow edging of crimson anteriorly and a 
wide margin of white or orange posteriorly (Forbes 
and Richardson 1920; Harlan and Speaker 1956; 
Clay 1962). Cook (1959) states that the opercular 
flap in males is broadly outlined in orange while in 
females the margin is white. Females are not as 
brilliantly coloured as males, their body colour 
tending towards olivaceous green, the flank spots 
being more dusky, the opercular spot being less 
contrasting and the fins transparent (Barney and 
Anson 1923; Becker 1983). As such, they more 
closely resemble immature males. The colouration 
of immatures is more subdued, and often includes 
rather distinct vertical bars and the lack of brownish 
spotting on the sides (Barney and Anson 1923; 
Pflieger 1975). Immatures are said to closely 
resemble Bluegill, Lepomis macrochirus (Smith 



1990 



NoLTiE: Status of the Orangespotted Sunfish 



71 



1979). Because of this, Canadian specimens may 
have previously gone unnoticed. 

Morphologically, the sexes also differ in outline, 
the angle between the skull and the dorsal profile at 
the nape being greater in males, yielding a more 
concave forehead than in females (Forbes and 
Richardson 1920). As such, reproductively active 
males were described as "high" by Cross (1950). 
Males also have longer ventral fins than do females 
(Forbes and Richardson 1920). 

Orangespotted Sunfish are peculiar among 
centrarchids in that they possess extremely large, 
sHt-like preopercular sensory pores within the lateral 
Une canals of the head (Moore 1957; Curd 1959; 
Branson and Moore 1962; Cross 1967; Becker 1983), 
the cranial and facial bones having been called 
"cavernous" by Ortenburger and Hubbs (1927). The 
two sensory pit depressions in the skull between the 
eyes are the largest among the sunfish, their widths 
being about equal to the distance between them 
(Pflieger 1975; Becker 1983). Also prominent are the 
openings along the free edge of the preopercule 
(Smith 1979; Trautman 1981; Becker 1983). These 
openings are much larger than the more rounded 



ones occurring in other centrarchids, their greater 
size possibly allowing improved detection of 
pressure waves in the obscured visual environment 
which prevails in the turbid waters of their typical 
habitats (Miller and Robison 1973). 

Morphologically, Canadian populations appear 
to deviate in some regards from southern ones. 
Holm and Coker (1981) provide fm ray counts for 
two Ontario capture locations, finding their 
numbers greater than in Kansas specimens 
examined by Cross (1967). Further counts of this 
kind (Jordan and Evermann 1902; Forbes and 
Richardson 1920; Eddy and Surber 1947; Cook 
1959; Eddy and Underbill 1974; Pflieger 1975; 
Becker 1983) also bear out this trend. These 
differences may be evidence that Canadian 
populations arose long ago and have since 
reproduced in isolation. Formal studies of meristic 
and morphological variation across the species' 
range would provide a useful test of this possibility. 

Distribution 

The Orangespotted Sunfish is endemic to North 
America, its distribution being restricted almost 




J 
100° 75' 

Figure 2. North American distribution of Lepomis humilis (adapted from Lee 1980). 



72 



The Canadian Field-Naturalist 



Vol. 104 



exclusively to the east-central United States, 
primarily in the Mississippi Valley (Figure 2). Its 
range extends north from Texas through Okla- 
homa, Colorado and the Dakotas, then east through 
southern Minnesota, the extreme southern portion 
of Wisconsin, Illinois and Indiana to western Ohio, 
and then south through Kentucky, Tennessee and 
Alabama. Within these boundaries, its distribution 
is continuous. However, it is most abundant in the 
west and southwest of its range (Hay 1894). Greene 
(1935) suggests that it underwent little post-glacial 
dispersal. Distribution details are provided 
elsewhere (Hay 1894; Meek 1895; Jordan and 
Evermann 1896, 1902; Forbes and Richardson 1920; 
Barney and Anson 1923; Greene 1935; Gerking 
1945; Eddy and Surber 1947; Moore and Buck 1953 
Eddy 1957; Moore 1957; Hubbs and Lagler 1958 
Cook 1959; Bailey and Allum 1962; Clay 1962 
Nelson and Gerking 1968; Smith-Vaniz 1968 
Pflieger 1971, 1975; Miller and Robison 1973; Buss 
1974; Eddy and Underbill 1974; Cross and Collins 
1975; McClane 1978; Smith 1979; Lee 1980; 
Trautman 1981; Becker 1983; Cooper 1983). 

Trautman (1981) concluded that the species' 
native range was restricted to the Mississippi 
drainage and hypothesized that this confinement 
was first broken around 1929 when specimens were 
discovered in Ohio's Lake St. Marys (the lake 
divides the Wabash/ Ohio/ Mississippi system and 
the St. Marys/ Maumee/ Lake Erie drainage). 
Trautman (1981) speculated that their presence in 
Lake St. Marys may have resulted from human 
activity, some individuals having been transported 
over the levee/ spillway at its western end. According 
to Trautman (1981) the species then expanded 
westwards. Portage River and Sandusky River 
specimens being taken in 1949. Immediately north 
of the mouths of these rivers lies Ohio's Bass Island 
chain, and specimens were taken there from South 
Bass Island in 1952. White et al. (1975) reported 
finding Orangespotted Sunfish in a Lake Erie 
tributary near Cleveland, Ohio, in 1971-72 and 
Trautman (1981) considered this a significant 
eastward extension of the range in Ohio. 

In contrast to the invasion scenario above, several 
lines of evidence point to Orangespotted Sunfish 
being native to the Lake Erie basin. Hubbs and 
Brown (1929) provide evidence of the species' 
presence in the Lake Erie basin upstream of Niagara 
Falls at a date earlier than the 1929 invasion 
suggested by Trautman ( 1 98 1 ). Nelson and Gerking 
(1968) considered the species to be a native of 
Indiana found in all major drainages (including 
Erie's) except those of Lake Michigan. Emery (1 976) 
also included the species among inhabitants of Lake 
Erie in the United States based on examination of 
unpublished commercial catch records, research 
vessel cruise reports, power plant entrainment 



studies and Great Lakes Fishery Commission 
reports, and classified the species as being 
uncommon in the lake but in stable populations. 
Likewise, Hocutt and Wiley (1986) suggested that 
these fish are native to tributaries of Lake Erie. 

In addition, Gerking (1945) states that Oranges- 
potted Sunfish were widely distributed in Indiana's 
downstream portion of the Maumee River as of 
1945, whereas Trautman (1981) suggests that the 
species only became established in the upstream 
portion of the Maumee in Ohio after 1929. 
Together, one might interpret these two reports as 
indicating a rapid and extensive invasion of the 
Maumee system. However, this seems unlikely given 
the time elapsed, suggesting instead that Oranges- 
potted Sunfish were native to the Maumee and thus 
to the Lake Erie watershed. 

Finally, Wise (1980) reported 1963 finds of 
Orangespotted Sunfish in Racoon Creek, a 
tributary of Ohio's Licking/ Muskingum River 
system which empties into Lake Erie at Lorain via 
the Black/ Kilbuck rivers. This evidence of the 
occurrence of Orangespotted Sunfish in eastern 
Ohio is significant in that it predates White et al.'s 
(1975) report from near Cleveland which Trautman 
(1981) used in postulating his range extension 
scenario. Instead, this early eastern Ohio find lends 
further support to the species being native to 
tributaries of Lake Erie. 

Despite the above, the species' presence in 
Canada probably does reflect some northward 
range expansion. Given its uncommon occurrence 
in Lake Erie (Emery 1976) it is not surprising that 
the species has only recently been discovered in 
Canadian waters. It may have existed in the latter 
for some time, as morphological differences have 
been noted (Holm and Coker 1981) between 
Canadian and United States populations. 

In Canada, populations of Orangespotted 
Sunfish have been discovered at six different 
southwestern Ontario localities (Table 1, Figure 3). 
The most southerly, from Pelee Island (Table 1), 
occurs ca. 24 km northeast of South Bass Island. 
Cedar Creek, on the Lake Erie north shore ca. 25 km 
north of Pelee Island, has produced specimens from 
four locations at various points along its lower 
reaches (Table 1). By their abundance here, these 
populations are apparently well-established (Holm 
and Coker 1981). A sixth site, Canada's most 
northern to date, occurs on the Canard River ca. 10 
km north of the Cedar Creek site furthest upstream. 
Noltie and Beletz (1984) provide evidence that it too 
supports a reproducing population. These Cana- 
dian populations may only be of localized 
abundance, since extensive Ontario Ministry of 
Natural Resources (OMNR) and National Muse- 
ums of Canada (NMC; now Canadian Museum of 
Nature) surveys conducted in 1979 and 1980 in the 



1990 



NoLTiE: Status of the Orangespotted Sunfish 



73 



Table I. Captures of Orangespotted Sunfish (Lepomis humilus) made in Canada to date. ROM = Royal Ontario 
Museum voucher specimen holdings. NMC = National Museums of Canada, Museum of Natural Sciences voucher- 
specimen holdings. 



Collection 



Capture 
Date 



Latitude 



Longitude 



Number 

Caught Location 



ROM 45577 15/05/84 



ROM 36442 11/06/i 



4I°48'42"N 82°39'31"W 



NMC 85-0529 14/07/83 42°0r00"N 82°47'00"W 



NMC 81-0055 11/06/80 42°01'30"N 82°49'18"W 



42°01'38"N 82°49'27"W 



ROM 36441 11/06/80 42°0r53"N 82°49'53"W 



Noltie & Beletz 21&23/06/83 42°07'24"N 82°50'51"W 

(1984) 



38* 



38* 



inside eastern breakwater, North 
Bay, Pelee Island, near Scudder 

Cedar Creek mouth at Lake Erie, 
Gosfield South Township, Essex 
County 

Cedar Creek, 0.5 km S of Arner, 
Colchester South and Gosfield 
South Townships, Essex County 

Cedar Creek, Colchester South 
Township, Essex County 

Cedar Creek, W of Arner, 
Colchester South Township, Essex 
County 

Canard River, 3 km SW of Gesto, 
Colchester North Township, Essex 
County 



*indicates two collections whose catch together totalled this number. 



immediate vicinity of these capture locations failed to 
locate Lepomis humilis specimens (G. Duckworth, 
OMNR Chatham, Ontario; personal communica- 
tion). Canadian Orangespotted Sunfish habitat may 
also differ seasonally, the populations sampled on 1 1 
June 1980 by Holm and Coker (1981) having been 
resampled by me on 15 July 1981 without success. 
Failure to encounter Lepomis humilis populations 
elsewhere in the Lake Erie and Ontario watersheds, 
despite widespread and vigorous sampUng by the 
OMNR, suggests that the restricted range indicated 
in Table 1 is relatively accurate. 

Protection 

The Orangespotted Sunfish is not specifically 
protected in Canada. Provisions of the federal 
Fisheries Act offer some general protection from 
potential habitat degradation, as do provincial and 
municipal statutes of this nature. Ontario fishing 
regulations permit the taking of this species year 
round. No size or possession limits apply. 

Population Sizes and Trends 

Estimates of population sizes and trends have 
not been established for the Orangespotted 
Sunfish in Canada. Although present populations 
appear small and narrowly distributed, trends in 
abundance and distribution elsewhere {see below) 
suggest that their range and numbers in Canada 
are likely to continue growing. 



Several reports propose that man has acted 
directly to increase the species' distribution. A 
more widespread occurrence in South Dakota west 
of the Missouri River suggests that inadvertent 
plantings with other species have occurred (Bailey 
and AUum 1962). Smith-Vaniz (1968) states that 
their presence in coastal drainage areas of 
Alabama east of the Mississippi and in a Georgia 
pond have similarly resulted from recent plantings. 
Captures have been made in a stocked Colorado 
reservoir (Gregory et al. 1979; Powell 1972, 1973). 
Farm pond introductions have extended its range 
within Ohio also (Trautman 1981). Thus, Lee 

(1980) contends that the general eastern and 
western spread of the species has resulted from 
unintentional introductions. 

Habitat alteration has also favoured range 
expansion in this species. More widespread 
distributions in Indiana have accompanied 
agricultural practices which transformed clear 
prairie-type streams to turbid plains-type ones 
(Gerking 1 945). Buth ( 1 974) and Smith ( 1 979) found 
the species more abundant and widespread after 
long-term increases in siltation in Illinois. Trautman 

(1981) relates the invasion of the species into central 
Ohio to increased stream turbidity and siltation of 
stream bottoms caused by clearing and tilling. 
Population sizes increased when small Ohio streams 
were channelized (Carline and Klosiewski 1985). 
Greater centrarchid abundance, with Lepomis 
humilis comprising a large portion, also resulted 



74 



The Canadian Field-Naturalist 



Vol. 104 



^g 


) 


GREAT LAKES BASIN 












so so tOUM 


jX 


S^';.^-,^ 






x^" 








^^^-nC^y-^^ 


\ ^.^-^^--T 


^'^-^ -— _^ '■^^ 




L 






STl**fNa 


-rLcc{] 


hi 


(5^-4( ^5: 


^M 


/ 


' ( 


1 / J <~S<^ 


'Jly ^ 


{ 


r \ 


\L/ \ 




' 


^^ 







Figure 3. Canadian (Ontario) distribution of the Orangespotted Sunfish. 



from installing floating tire breakwaters in an 
Oklahoma reservoir (Clady et al. 1979). Similar 
habitat changes may be expected to favour 
Canadian populations. 

In contrast, human influences have also 
affected some populations adversely. Larimore 
and Smith (1963) report that the species' 
abundance and distribution shrank in parts of 
Illinois with stream changes caused by man. In 
Kansas, the species' distribution also failed to 
increase despite lake construction and stocking 
programs (Cross 1967). Populations have 
declined in areas of preferred habitat in Illinois, 
despite changes tending to produce more habitat 
apparently favourable to the species (Smith 
1968).' Reduced abundance in Illinois has 
accompanied draining of the prairie wetlands 
(Smith 1979). Isolation from a feeder stream, 
dredging, and stocking with Pike {Esox lucius) 
and Largemouth Bass (Micropterus salmoides) 
completely decimated one Iowa pond population 
(Carlander 1978). In Colorado, changes since 
European settlement have caused the species' 
range in the South Platte system to shrink 
sufficiently for it to be designated rare (Propst 
and Carlson 1986), although in none of the 
American states are populations considered 
threatened (Miller 1972; Johnson 1987). 
Obviously, large-scale habitat changes like those 
above are likely to impact negatively on 
Canadian populations. 



Habitat 

Many authors provide information on the 
general habitat preference of the Orangespotted 
Sunfish. In the lower Missouri River basin, the 
species was said to reach local abundance in sandy 
streams (Jordan and Evermann 1902). Tolerance 
of a wide range in pH, and rapid changes thereof 
was shown by Wiebe (1931), who observed 
spawning in waters as high as pH 9.3. Inhabitation 
of turbid waters in the Great Plains region may 
have been responsible for the evolution of the 
species' large lateral line canals (Moore 1956). 
Orangespotted Sunfish reportedly frequent turbid 
lentic, and sluggish lotic waters, often to the 
exclusion of other sunfishes (Branson and Moore 
1962). Miller (1963) considered the species tolerant 
of mud and silt in ponds and streams. Where it 
occurs outside of Great Plains areas, stream 
habitats resembling those of the Plains are chosen 
(Metcalf 1966). Lee (1980) describes its habitat as 
being quiet streams and vegetated lakes or ponds. 
Cooper (1983) lists it as an inhabitant of turbid, 
silty streams and ponds. Hocutt and Wiley (1986: 
1 69) list the species as a dweller in both upland and 
lowland streams and big rivers. 

Despite the generalizations above, Orangespot- 
ted Sunfish habitat varies considerably throughout 
the range. An effort has been made to draw 
together the literature pertaining to this subject 
and a summary is presented in Table 2. Since co- 
occurring species are also important components 



1990 



NoLTiE: Status of the Orangespotted Sunfish 



75 



Table 2. Summary of literature references dealing with the habitat preferences of Lepomis humilis, and their species 
associates age and growth rates. 



Area 



Habitat 



Species Associates 



Age & Growth 



Alabama Tarzwell (1945) 



Colorado Powell (1973), Propst and Carlson 
(1986) 

Illinois Forbes and Richardson (1920), 

Thompson and Hunt (1930), 
O'Donnell (1935), Brown and 
Thompson (1937), Whitacre (1952), 
Stegman (1958, 1959), Larimore and 
Smith (1963), Smith (1968, 1979) 

Indiana Gerking ( 1 945) 

Iowa Meek (1895), Barney and Anson 

(1923), Harlan and Speaker (1956), 
Mitchell etal. (1980) 



Kansas Cross (1954, 1967), Minckley (1956, 
1959), Prophet (1957), Metcalf 
(1959, 1966), Deacon (1961), Deacon 
and Metcalf (1961), Gash (1967), 
Gash and Gash (1973), Cross and 
Collins (1975) 

Kentucky Carter ( 1 954), Clay ( 1 962) 

Louisiana Barney and Anson (1923), Carver 
(1967) 

Minnesota Eddy and Surber (1947), Eddy and 
Underbill (1974) 

Mississippi Cook (1959), Font et al. (1984) 

Missouri Pflieger (1971, 1975) 

Nebraska Meek (1895), Canfield and Wiebe 
(1931) 



Ohio 



Wise (1980), Trautman (1981) 



Oklahoma Moore and Paden (1950), Cross and 
Moore (1952), Ward (1953), Finnell 
(1954), Elkin (1955), Jenkins et al. 
(1955), Gould and Irwin (1965), 
Branson (1967), Bross (1969), Miller 
and Robison (1973), Margraf and 
Plitt(1982) 



South 
Dakota 



Wisconsin 



Greene (1935), UMRCC (1948b), 
Christenson and Smith (1965), 
Becker (1983) 



Tarzwell (1945), Alabama 
Department of Conservation 
(1958), Smith-Vaniz( 1968) 

Barnhart 1955 



Forbes and Richardson (1920), 
Cahn (1927), Bennett (1943), 
Upper Mississippi River 
Conservation Commission* 
(1948a), Stegman (1958, 1959), 
Lopinot (1967, 1968), Buth (1974) 

Whitaker etal. (1977) 

Meek (1985) UMRCC (1948a) 



Moore and Buck (1953), Minckley 
(1956, 1959), Schelske( 1957), 
Metcalf (1949, 1966), Cross (1954, 
1967), Deacon and Metcalf (1961), 
Gash (1967), Gash and Gash 
(1973) 

Carter (1954) 

Barney and Anson (1923) 



Clark (1960), Hoke et al. (1979), 
Laskowski-Hoke et al. (1982), 
Carline and Klosiewski (1985) 

Hubbs and Ortenburger (1929), 
Cross (1950), Moore and Paden 
(1950), Thompson (1950), Buck 
and Cross (1951), Cross and 
Moore (1952), Moore and Buck 
(1953), Branson (1967), Orth 
(1980) 

Shields (1955), Bailey and Hubbs 
(1962), Wahl and Applegate (1981) 

Cahn (1927), UMRCC* (1948b), 
Christenson (1957), Christian and 
Smith (1965) 



Swingle (1965) 



Barnhart 1955, Gregory et al. 
(1970), Powell (1972, 1973) 

Whitacre (1952), Lopinot (1958), 
Stegman (1958), Starrett and Fritz 
(1965) 



Barney and Anson (1923), 
Carlander (1949), Carlander and 
Parsons (1949), Carlander and 
Parsons (1949), Harlan and 
Speaker (1956) 

Minckley (1959) 



Barney and Anson (1923), Carver 
(1967) 



Cook (1959) 



Clark (1960), Trautman (1981) 



Cross (1950), Buck and Cross 
(1951), Jenkins (1953), Finnell 
(1954), Elkin (1955), Jenkins et al. 
(1955), Jenkins and Finnell (1957), 
Gould and Irwin (1965), Whiteside 
(1967), Whiteside and Carter (1973) 



Shields (1955) 



Meehean (1932), Christenson 
(1957), Christenson and Smith 
(1965), Becker (1983) 



"UMRCC = Upper Mississippi River Conservation Commission. 



76 



The Canadian Field-Naturalist 



Vol. 104 



of habitats, studies documenting these are also 
noted. 

In Canada, the Cedar Creek populations (Table 
1) were found in shallow muddy waters with 
bottoms of silt, clay and detritus (Holm and Coker 
1981). Canard River specimens came from like 
habitat, the sample site being shallow, slow- 
flowing, turbid, and free of aquatic macrophytes, 
with a primarily clay-silt bottom (Noltie and Beletz 
1984). Species co-occurring there were similar to 
many of the species suites referred to elsewhere 
(Table 2) for American populations. As such, 
Canadian populations seem to have maintained 
the general habitat preferences they exhibit in the 
United States. The Lake Erie population at Pelee 
Island is somewhat anomalous, however, since 
large lakes appear to be rarely inhabited by this 
species. 

General Biology 

Age and Growth: Orangespotted Sunfish are 
usually said to reach 9 to 10 cm in length (Jordan 
and Evermann 1896, 1902; Richardson 1904; 
Forbes and Richardson 1920; Eddy and Surber 
1947; Ward 1953; Eddy 1957; Clay 1962; Cross 
1967; Buss 1974; Eddy and Underbill 1974; Cross 
and Collins 1975; McClane 1978). Lengths of 5 to 
7.5 cm are more typical (Hay 1894; Eddy and 
Underbill 1974), but individuals 12 to 16 cm long 
have also been recorded (Barnhart 1955; Jenkins et 
al. 1955; Lopinot 1958; Swingle 1965; Carver 
1967), these being considered exceptional. Seven 
years is the maximum recorded age, although 
individuals in most populations fail to live past 
their fourth year, and growth is not slowed initially 
by the attainment of sexual maturity (Barney and 
Anson 1923). Some Oklahoma reservoir popula- 
tions are apparently annuals (Jenkins 1953; Finnell 
1954; Elkin 1955). Carlander (1977) provides a 
summary of growth and condition in the species. 
Recorded standing crop estimates for the species 
are also summarized by Carlander (1955) and 
Whiteside and Carter (1973). Various studies of 
age and growth in this species have been 
conducted, and the references are summarized in 
Table 2 by state. These indicate that the lifespan of 
southern individuals is often shorter than in the 
north, and that maturity at age classes II and III is 
more typical. Southern specimens generally grow 
more rapidly, due likely to the longer feeding and 
growing seasons and earlier breeding period. 
Growth occurs primarily in late spring and early 
summer (Barney and Anson 1923; Jenkins et al. 
1955; Shields 1955). Trautman (1981) and Becker 
(1983) document the sizes of young of the years in 
Ohio and Wisconsin, respectively. 

In Canada, only two populations have been 
sampled for age and growth. Cedar Creek 



individuals ranged between 36 and 97 mm total 
length (TL). Females of 51 to 58 mm TL were 
gravid, and the two largest males (83 and 97 mm TL) 
were older males with enlarged ear flaps. Smaller 
specimens there were males and/ or juveniles (Holm 
and Coker 1981). Canard River specimens ranged 
from 45 to 107 mm TL and age classes II to IV, 
males being the largest among the collected 
specimens. A length-weight regression for this 
sample was calculated by Noltie and Beletz (1984). 
Together, data from these two studies indicate that 
Canadian Lepomis humilis are of sizes and occur in 
populations with age structures similar to those 
described from elsewhere in the range. 

Diet: Information on the diet of Canadian 
Orangespotted Sunfish does not exist. However, 
food studies have been made on natural 
populations from further south (Barney and 
Anson 1923; Cahn 1927; Hildebrand and Towers 
1927; Rice 1941; Lagler and Ricker 1942; Clark 
1943; Harrison 1950; Ward 1953; Kutkuhn 1955 
Stegman 1958, 1959; Wenke 1965; Whitaker 1975 
Gaudet personal communication: in Becker 1983) 
These indicate that aquatic insects at various life 
stages and crustaceans compose most of the diet. 
Midges, mayflies, caddis flies and water boatmen 
dominate the aquatic insect component, while 
amphipods, cladocera, copepods, isopods and 
ostracods constitute the crustacean portion. Water 
mites, bryozoans, small fish, and plant material are 
also important. Kutkuhn's (1955) analysis of 
specimens from a dredged lake indicated that the 
insects consumed originate primarily in shallow 
water. Clark (1943) makes a similar conclusion 
based on diet studies from other lake specimens. 
Cross and Collins (1975) also consider their diet in 
Kansas to be mostly insects. The species' long, 
pointed and slender gill rakers likely facilitate 
taking such foods (Jordan and Evermann 1902; 
Forbes and Richardson 1920; Eddy and Surber 
1947; Eddy 1957; Clay 1962; Cross 1967; Eddy and 
Underbill 1974). Larger food items tend to be 
consumed as individuals surpass the yearling stage 
(Kutkuhn 1955). Larger Lepomis humilis also eat 
smaller ones on occasion (Barney and Anson 1923; 
Lagler and Ricker 1942). The species has been 
classified as one having a food chain of 
intermediate length (Carlander 1955). Intra- 
specific competition for such food resources can be 
intense, large year classes decreasing in mean 
weight per individual (Orth 1980). 

In the laboratory, various aquarium studies 
indicate that specimens fare well on diets of 
daphnia and other live foods (Luce 1938; Harlan 
and Speaker 1956), live Daphnia magna (Gould 
and Irwin 1965), or dried commercial flake food, 
Daphnia. Chironomus larvae, and earthworms 
(Dennis 1970; Powell 1972). 



1990 



NoLTiE: Status of the Orangespotted Sunfish 



77 



Behaviour: Miller (1963) and Miller and 
Robison (1973) considered the Orangespotted 
Sunfish more gregarious than other Lepomis 
species such as the Green Sunfish. Their feeding 
was described as voracious by both Branson and 
Moore (1962) and Gould and Irwin (1965). 
Although pugnacious (Branson and Moore 1962), 
nesting males may frighten sufficiently to 
temporarily vacate their nests (Cross 1967). 
Miller's (1963) classic work describes comfort 
movements, and feeding, sleep, locomotory, 
agonistic, social, nesting, courtship and spawning 
behaviours in the species. Captive specimens adapt 
easily to holding conditions (Gould 1962; Gould 
and Irwin 1965; Irwin 1965), although Miller 
(1963) found them difficult to acclimatize and the 
species quite timid. Aspects of their agonistic 
behaviour and social organization have been 
examined in the laboratory (Dennis 1970; Powell 
1972; Petersen 1979), common behaviours 
including approach, fin erection, bite, chase, 
opercular spread, tail beat, and avoid. Behavioural 
changes among males, primarily decreased levels 
of aggression, have been used effectively in 
bioassay work examining the sublethal effects of 
wastewaters (Shelford 1917; Petersen 1979; 
Petersen et al. 1986). Changes in iris colouration, 
abihty to retain equilibrium, feeding habits, 
exaggerated breathing movements, and irritability 
were also observed (Petersen 1979). There is some 
indication that individuals are more active in 
summer months than in winter (Petersen 1979). 

Reproduction: Little is known of the breeding 
biology of Orangespotted Sunfish in Canadian 
populations. Holm and Coker (1981) report that 
Cedar Creek females were gravid 1 1 June. Noltie 
and Beletz (1984) found sexually mature Canard 
River males of ages two through four years to have 
been greater than 55 mm TL. These had free- 
flowing milt when caught on 21 and 23 June 1983. 
No further information on Canadian populations 
exists. 

In southern populations, reproductive maturity 
is reached relatively early in life. Breeding must 
occur in the first year of life in the single age class 
populations reported from Oklahoma (Jenkins 
1953; Finnell 1954; Elkin 1955). Barney and Anson 
(1923) found that some Lepomis humilis young 
matured in their second year, but that the majority 
laid their first eggs at the beginning of their third 
year. The smallest gravid female they measured 
was only 3 cm in length. Cook (1950) stated that 
Mississippi individuals often spawn when only 6 
cm long, while the smallest reproductive individual 
in Whitacre's (1952) Illinois lake population was 37 
mm standard length (SL). Lee's (1980) statement 
that a minimum size of 48 mm is required for 
reproductive maturity thus appears in error. 



Information on the breeding season of the 
Orangespotted Sunfish is scattered and fragmen- 
tary. The following information is arranged in 
order of increasing latitude. In Louisiana, 
spawning begins in early April and continues until 
September, the critical onset temperature being 
18°C (Barney and Anson 1923). Well-developed 
young are seen as early as late April. In Mississippi, 
Hildebrand and Towers (1927) captured ripe 
specimens between June and September. In 
Oklahoma, breeding individuals were found in late 
May when temperatures reached 25° C, and young- 
of-the-year were taken by early July at 28° C (Cross 
1950). The height of the spawning season is thus in 
early spring in Oklahoma, but may extend to July 
or August (Ward 1953). Brood stock seeded in an 
Oklahoma pond in May spawned and produced 
seinable offspring by August of the same year 
(Gould and Irwin 1965). In Kansas, nesting was 
observed in late June and early July (Minckley 
1959). The peak in spawning activity occurs in 
Kansas in late May and June, water temperatures 
spanning 24° to 32° C (Cross 1967), In Missouri, 
Pflieger (1975) states that nesting occurs from late 
May into August. In Illinois, males and females in 
breeding colour were observed over new nests in 
late May in a marsh. Gravid females and brightly 
coloured males were seen as late as early July in an 
adjacent lake (Richardson 1913). Pairs of ripe 
males and females in breeding colouration were 
found in IlUnois in early June (Forbes and 
Richardson 1920). Cahn (1927) captured an 
Illinois male in full breeding colouration in early 
July. Young-of-the-year were taken in the latter 
part of June by Whitacre (1952). Gonad 
examinations also confirm a May and early June 
spawning season in Illinois (Stegman 1958, 1959). 
In Iowa, breeding begins in late May and ends in 
August (Barney and Anson 1923; Harlan and 
Speaker 1969). In Wisconsin, spawning takes place 
from May to July (Eddy and Surber 1947). Becker 
(1983) found that spawning occurred in Wisconsin 
from late May to August, beginning at water 
temperatures of about 18°C and continuing to 24° 
to 32° C. Lee (1980) suggests that nesting occurs 
from May to July in northern portions of the 
range, and from April to September in the south. 
The peak spawning period occurs in late April or 
early May, although stragglers extend this to early 
August (McClane 1978). Younger fish nest later in 
the season and geographical location and weather 
conditions also modify season length (Barney and 
Anson 1923). 

Orangespotted Sunfish are classified among the 
nest-spawning lithophilic fishes which guard their 
young (Balon 1975; Orth 1980). Males construct 
the nests (Buss 1974) which are roughly circular, 
are 3 to 4 cm deep, and are excavated by the males 



78 



The Canadian Field-Naturalist 



Vol. 104 



either pushing or sweeping the substrate into 
position (Barney and Anson 1923). As this 
sweeping motion occurs, males move from a 
horizontal to a vertical, head-up position, making 
little horizontal movement through the nest 
(Miller 1963; Keenleyside 1979). Nests are situated 
where sand or fine gravel can be exposed (Cross 
1967; Buss 1974), or are dug through mud and silt 
until sohd bottoms of gravel are reached (Barney 
and Anson 1923). Branson (1967) suggests that 
nests may also be constructed in rocks and gravel. 
Where only a mud bottom occurred, Louisiana 
males constructed no distinct nests (Barney and 
Anson 1923). 

Nests have been observed at depths of 46 cm in 
an Illinois marsh by Richardson (1913), at depths 
of 30 to 91 cm in Iowa where they were placed in 
areas of soft bottom and measured 15 to 18 cm in 
diameter (Barney and Anson 1923), and in water 
depths from 10 to 61 cm in Kansas (Cross 1967). 
Thus, nests are generally in water less than 90 cm 
deep, although occasionally spawning occurs at 
depths just sufficient to cover the backs of the 
spawners (McClane 1978). 

Males typically nest in clusters, the maximum 
recorded size apparently being a colony of nearly 
1000 which formed a band about a metre wide 
along a ca. 1 10 m bank in Iowa (Barney and Anson 
1923). Within these colonies, males defend 
individual territories 30 to 60 cm in diameter 
(Cross 1967). 

Barney and Anson's (1923) study of reproduc- 
tion is the most complete to date, and yields the 
following information. After completing his nest, a 
male attracts a female to it. Spawning then occurs 
in typical centrarchid fashion, the pair adopting a 
parallel position over the nest, whereupon eggs and 
sperm are released. The female then departs. The 
mating system is one of promiscuous polygamy. 
Males guard the eggs, which measure 1 mm in 
diameter and are adhesive. Fanning of the eggs by 
the parental male prevents siltation. Males also 
defend the nest territory aggressively. Darters, 
minnows, and adjacent males all prey on the nest 
contents. The egg stage lasts about five days at 
water temperatures of 18° to21°C. Newly hatched 
young are 1 cm in length. Females contain up to 
4700 eggs, fecundity varying with size and progress 
through the breeding season (Barney and Anson 
1923). 

Hildebrand and Towers (1927) report that one 
60 mm female contained nearly 1200 ripe eggs, 
double the number contained by two others of 
nearly the same length. A 10.8 cm female from 
Oklahoma bore about 5000 eggs (Ward 1953). 
Eddy and Surber(l943: in Brederand Rosen 1966) 
reported 15 000 to 58 000 eggs per fish in 
Wisconsin females, a figure which is not repeated 



in a subsequent edition of this book. Further 
information on fecundity is provided for Colorado 
females by Beckman (1952), and for Wisconsin 
females by Becker (1983). Only 25 to 300 of these 
eggs are laid through a spawning bout (Buss 1974; 
McClane 1978), these being deposited in numbers 
of between 5 to 15 on each tilt of the male and 
female (Miller 1963). The eggs are minute and 
nearly colourless, appearing as grains of clear 
quartz sand, and adhere to the nest substrate 
(Cross 1967). The adhesive, amber-coloured eggs 
produced by Wisconsin females measured 0.5 mm 
in diameter (Becker 1983). Minnows are acknowl- 
edged egg predators, and swarm into temporarily 
deserted nests (Cross 1967). 

The young and their pigmentation are described 
in Auer (1982), yolk-sac larvae measuring 5.3 mm 
TL, larvae 7.9 mm TL. Red Shiners (Notropis 
lutrensis) and Redfin Shiners {Notropis umbrati- 
lis) have been known to spawn over nests of this 
species in Missouri (Pflieger 1975). The grunting 
sounds used by Lepomis humilis males to attract 
conspecifics during courtship and how these 
sounds differ from those of other Lepomis species 
are documented by Gerald (1971). 

Hybridization: Hybrids frequently occur 
between Orangespotted Sunfish and Bluegill 
(O'Donnell 1935; Luce 1938; Cross and Moore 
1952; Stegman 1958; Breder and Rosen 1966; 
Branson 1967; Mitchell et al. 1980), Green Sunfish 
(Hubbs and Ortenburger 1929; Hubbs and Hubbs 
1933; Breder 1936; Luce 1938; Minckley 1959; 
Bailey and Allum 1962; Zach 1968; Bross 1969; 
Trautman 1981), Longear Sunfish, Lepomis 
megalotis (O'Donnell 1935; Cross 1950), Pump- 
kinseed, Lepomis gibbosus {0'T>onne.\\ 1935; Luce 
1938; Upper Mississippi River Conservation 
Committee 1948b; Trautman 1981), and Redear 
Sunfish, Lepomis microlophus (Trautman 1981), 
and with the Warmouth, Lepomis gulosus 
(Whitmore 1986). Slastenenko (1957) and 
Schwartz (1972) list further sources reporting these 
hybrid combinations. Such hybrids reach greater 
size then do purebred L. humilis specimens (Eddy 
and Underbill 1974). 

This frequent hybridization may occur because 
Orangespotted Sunfish occasionally breed in areas 
used by other Lepomis species (Minckley 1959; 
Gerald 1971; Eddy and Underbill 1974), increasing 
opportunities for inter-specific pair formation. 
Alternatively, Branson (1967) suggested that such 
hybrids form because Orangespotted Sunfish are 
occasionally forced to spawn with congeners with 
which they become trapped when small, sluggish 
streams are reduced to intermittent pools for long 
periods. The use of species-specific courtship 
sounds (Cierald 1971) may aid in reducing such 
pairings, however. 



1990 



NoLTiE: Status of the Orangespotted Sunfish 



79 



Trautman (1981) states that in cases of Lepomis 
humilis range expansion, the leading front is 
invariably associated with frequent inter-specific 
hybridization, immigrants in small numbers being 
forced to mate with members of other species. 
However, once Lepomis humilis populations are 
well established, hybrids are less frequently 
encountered. Accordingly, Holm and Coker 
(1981) found that hybrids with Pumpkinseed 
constituted more than 10% of their collections 
from Canadian populations. Trends in hybrid 
numbers should be watched should Canadian 
populations prove non-ephemeral. 

Parasites: Because of its recent occurrence in 
Ontario waters, a listing of parasites for Canadian 
populations is not provided by Margolis and 
Arthur (1979). Hoffman (1967) supplies a 
summary of the known parasites of the species 
across North America, but does not include the 
following. Coker et al. (1922) report that 
Orangespotted Sunfish are infected naturally by 
glochidia of the Yellow Sand Shell, Lampsilis 
anodontoides, a freshwater clam. Cross (1960) 
reports that in Ohio populations Lernea carassii 
infects the species. McDaniel and Bailey (1966) 
report that cestodes, nematodes, and various 
trematodes parasitized Lepomis humilis specimens 
from Oklahoma. Gash (1967) and Gash and Gash 
(1973) list trematode, cestode, and nematode 
parasites from Kansas samples, and found greater 
numbers of some species in larger fish indicating 
that infection may occur on a cumulative basis. 
The crustacean Lernaea cyprinacea heavily 
parasitized some specimens in Indiana (Whitaker 
and Schleuter 1975), and monogenetic trematodes 
infested Lepomis humilis specimens from 
Oklahoma (Petersen 1979). Crites et al. (1984) 
found the species infected heavily with Neascus- 
type trematode metacercaria, and natural 
metacercarial infections by the parasite Phagicola 
nana occurred in collections from brackish 
estuaries of the Mississippi Gulf Coast (Font et al. 
1984). Larger fish were more heavily infected and 
the distribution of parasites through the body 
greater. 

Concerning resistance to parasitism, Anson and 
Howard {in: Barney and Anson 1923) state that the 
species is the only centrarchid which resisted 
infection by glochidia of the clam Lampsilis 
luteola. The gall bladder parasite Chloromyxum 
trijugum also failed to infect Orangespotted 
Sunfish or hybrids with Bluegill (Mitchell et al. 
1980). 

Limiting Factors 

Canadian populations of Orangespotted 
Sunfish may be limited by competition with other 
small native centrarchids. Competition with 



crappies (Pomoxis sp.), Bluegill and young bass 
{Micropterus sp.) for food was considered likely by 
Cross (1967). Competition with Green Sunfish was 
suggested as being partially responsible for the 
scarcity of Orangespotted Sunfish in Colorado 
(Propst and Carlson 1986). Because such 
competing species are widely distributed across 
southern Ontario, Lepomis humilis may be 
prevented from flourishing. In contrast, however, 
differences in feeding structures have also been 
suggested to reduce competition between Lepomis 
humilis and crappies (Forbes and Richardson 
1920), and their ability to outcompete Bluegill was 
expected to herald increased abundance in a 
polluted Oklahoma stream (Cross 1950). 

Predation by other species may also impose 
limitations on the species in Ontario systems. As 
outlined later, species of native southern Ontario 
fishes such as Largemouth Bass and Muskellunge 
are known predators of Orangespotted Sunfish. 
These may also mediate Lepomis humilis' 
population growth. 

The continued degradation of southern Ontario 
streams may also limit their spread and numerical 
increase. Given that the species is a typical 
inhabitant of turbid waters, continued field 
draining, stream channelization, and increased 
siltation and turbidity appear initially favourable to 
the species. However, Smith (1968) found that even 
where such changes occurred, Lepomis humilis was 
decimated in Illinois rivers. Limits to which habitat 
changes benefit the species thus exist. 

Concerning pollution, Orangespotted Sunfish 
were considered the most tolerant of the species in 
an Oklahoma stream (Cross 1950). In particular, 
the species' tolerance for oil refinery wastewaters is 
considered good (Gould 1962; Gould and Irwin 
1965; Irwin 1965), perhaps important given the size 
of southwestern Ontario's petrochemical industry. 
Despite such resilience, many fractions of oil 
refinery waste are lethal to Orangespotted Sunfish, 
especially among smaller individuals (Shelford 
1917). Early mortality due to oil pollution was 
suggested by the findings of Jenkins and Finnell 
(1957) also. Such exposure alters their agonostic 
behaviour (Petersen 1979; Petersen et al. 1986), 
and since territoriality is important during nesting 
in this species, deleterious effects on breeding 
populations may be expected. Zinc and cadmium 
concentrations in a small sample were high relative 
to a variety of species examined by Murphy et al. 
(1978), suggesting that the accumulation of 
environmental contaminants might limit Lepomis 
humilis. Indiviuals also accumulate organochlo- 
rine, heavy metal, and other elemental contami- 
nants (Lowe aet al. 1985; Schmitt et al. 1985). 
Clark (1960) also refers to the occurrence of 
thyroid hyperplasia in Ohio specimens. 



80 



The Canadian Field-Naturalist 



Vol. 104 



Special Significance of the Species 

The Orangespotted Sunfish belongs to the most 
species rich of the centrarchid genera (Avise and 
Smith 1977). Historically, the genus Lepomis iirst 
appears at the Miocene-Pliocene boundary (Miller 
1965). Fossil jaw bones of Lepomis humilis 
containing its characteristically enlarged sensory 
canal pores date the species back approximately 
1.2 million years to the -early Pleistocene (Bennett 
1977; Bennett 1979). Branson and Moore's (1962) 
consideration of centrarchid lateral line systems 
provides evidence of the species' taxonomic status 
within the family, differences in their lateral Une 
pore shape once having been responsible for their 
placement in the genus Allotis (Ortenburger and 
Hubbs 1927). The congener closest to them is the 
Bluegill (Bailey 1938; Avise and Smith 1974, 1977; 
Whitmore 1986), which may explain the similari- 
ties in their behaviour (Miller 1963) and the 
frequency of hybrids between the species (see 
above). Chromosome numbers and morphology in 
Lepomis humilis suggest that the species is an 
offshoot of the main centrarchid hne (Roberts 
1964a). Whereas all other Lepomis species have 48 
chromosomes (Roberts 1964a), the diploid count 
in Lepomis humilis is 46 (Roberts 1964a,b, 1967; 
Becker 1983). Existing genetic variability estimates 
(Avise and Smith 1974, 1977; Avise 1977; Avise et 
al. 1977) form the basis upon which evidence of 
founder effects in Canadian populations could be 
examined. 

From an aesthetic standpoint, Orangespotted 
Sunfish are unique among Ontario's fish species. 
Spawning males are said to be the most strikingly 
colourful of all sunfishes (Hay 1894; Jordan and 
Evermann 1902; Forbes and Richardson 1920; 
Cahn 1927; Hildebrand and Towers 1927; Eddy 
and Underbill 1974; Cross and Colhns 1975). 
During the breeding season; "the males are so 
brilliantly coloured they appear to be more 
artificial than real" (Harlan and Speaker 1956) and 
because of this they are attractive as aquarium 
fishes. 

Larger specimens may be occasionally kept 
when angled (Starrett and Fritz 1965), and are 
considered a valuable pan fish (Evermann and Cox 
1896). They apparently bite readily (Jordan 
Evermann 1902; Harlan and Speaker 1956), 
worms and grasshoppers being effective baits 
(Cross 1967; Eddy and Underbill 1974). Because of 
their small size, most anglers value them little as 
sport fish (Hay 1894; Jordan and Evermann 1902; 
Jenkins et al. 1955; Harlan and Speaker 1956; 
Cross 1967; Buss 1974; Eddy and Underbill 1974; 
Cooper 1983). Hence their species name, meaning 
"humble" (Jordan and Evermann 1896). 

Orangespotted Sunfish are utilized by other 
species in varying degrees as a forage species. 



Barney and Anson (1923) found that their numbers 
fell substantially in late summer due to predation 
by other fishes. Thus, one value of the species lies in 
its use as forage for other fishes of angling quality, 
especially in cultures of Largemouth Bass. The 
potential for their control of mosquitoes is also 
discussed by these authors. Lagler and Ricker 
(1942) found that Yellow Bass {Morone mississip- 
piensis) ate Orangespotted Sunfish. Cross (1967) 
considers that where abundant, they may also 
provide significant forage for larger centrarchids. 
Krska and Applegate (1982) found Lepomis 
humilis to be an important food resource of young 
Muskellunge. However, Buck and Cross (1951) 
declared the species to be "generally considered 
detrimental", and that specimens were rarely 
found in game fish gut contents, perhaps due to 
their long dorsal spines. 

Orangespotted Sunfish may also impact 
negatively on other more desirable species. Luce 
(1933) considered it a "weed" since it consumes 
food likely to be useful to larger fishes. Lagler and 
Ricker (1942) as well as Clark (1943, 1960) and 
Cross (1967) suggest them to be undesirable in 
some locations because of their competition for 
food with more valuable species. Bennett (1943) 
states emphatically that the species "never grows to 
a desirable size and should never be stocked in 
artificial lakes", given that large populations had 
crowded out other centrarchids in some situations. 
Because of their diminutive size and competition 
with Bluegill, Shields (1955) also considered them 
undesirable. 

Clark (1960) states that Lepomis humilis also 
compete for nesting areas with Bluegill. Where 
sympatric, competition with the Green Sunfish 
may also occur (Trautman 1981). In contrast, 
Forbes and Richardson (1920) suggest that their 
competition with other centrarchids is minimal 
due to differences in their feeding structures. 

Also of significance is that the species has been 
used as an experimental subject in various toxicity 
investigations (Shelford 1917; Gould 1962; Gould 
and Irwin 1965; Petersen 1979; Petersen et al. 
1986), and has shown itself to be an excellent test 
and laboratory animal. 

Evaluation 

Orangespotted Sunfish populations located in 
Ontario appear to be indigenous, are extremely 
few, and are restricted in distribution. Captures 
have shown them to be generally in low 
abundance, although evidence points to some 
populations being self-perpetuating. The known 
distribution pattern and high frequency of hybrids 
in some catches suggests that Canadian popula- 
tions represent the leading edge of a northward 
range extension by the species. Because their 



1990 



NoLTiE: Status of the Orangespotted Sunfish 



Canadian distribution is very restricted and on the 
fringe of the species' range, and their numbers are 
low but do not appear to be in immediate peril, a 
designation of vulnerable would appear to be 
appropriate at this time. 

Acknowledgments 

Information on Canadian capture locations was 
graciously supplied by G. E. Gale and G. A. 
Goodchild of the Ontario Ministry of Natural 
Resources, D. E. McAllister and B. W. Coad of 
the National Museum of Natural Science, (now 
Canadian Museum of Nature) and E. J. Grossman 
of the Royal Ontario Museum. A. Kutas 
researched much of the literature. Support for the 
project was provided under contract to the Ontario 
Ministry of Natural Resources. Reviews by E. J. 
Grossman, D. E. McAllister, J. S. Nelson and 
A. E. Peden improved the manuscript. 

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the reproductive guild concept. Proceedings of the 
Oklahoma Academy of Science 60: 10 17. 

Petersen, J. C. 1979. Sublethal effects of biologically 
treated petroleum refinery wastewaters on agonostic 
behavior of male orangespotted sunfish, Lepomis 
humilis (Girard). M.Sc. thesis, Oklahoma State 
University, Stillwater, Oklahoma. 

Peterson. J. C, S. L. Burks, and R.J. Miller. 1986. 
Sublethal effects of biologically treated petroleum 
refinery wastewaters on agonistic behavior of male 
orangespotted sunfish, lepomis humilis (Girard). 
Environmental Joxicology and Chemistry 5(5): 
463 471. 

Pnieger,W. L. 1971. A distributional study of Missouri 
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Natural History 20(3): 225 570. 



1990 



NoLTiE: Status of the Orangespotted Sunfish 



85 



Pflieger, W. L. 1975. The fishes of Missouri. Missouri 
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Missouri. 

Powell, L. E. 1972. Factors influencing agonistic 
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Powell, T. G. 1973. Effect of northern pike introduc- 
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Rice, L. A. 1941. The food of six Reelfoot Lake fishes in 
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Richardson, R. E. 1904. Areviewofthesunfishesofthe 
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Richardson, R. E. 1913. Observations on the breeding 
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Roberts, F. L. 1964a. A chromosome study of twenty 
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Roberts, F. L. 1964b. A chromosome study of the 
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Roberts, F. L. 1967. Chromosome cytology of the 
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Whitaker, J. O., Jr. 1975. Foods of some fishes from the 
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Accepted 10 October 1989 



Status of the Bigmouth Buffalo, Ictiobus cyprinellus, in Canada* 
Cheryl D. Goodchild 

2168 Harcourt Crescent, Mississauga, Ontario L4Y IWl 

Goodchild, Cheryl D. 1990. Status of the Bigmouth Buffalo, Ictiobus cyprinellus, in Canada. Canadian Field- 
Naturalist 104(1): 87-97. 

The Bigmouth Buffalo, Ictiobus cyprinellus, is rare in Canada where its distribution is disjunct and exceedingly 
limited. In Saskatchewan, populations in some streams are large enough to support a limited commercial fishery but 
elsewhere in Manitoba and Ontario, populations are extremely small and tenuous judging from the very few specimens 
that have been collected. In contrast, the species is widespread and abundant throughout the Mississippi River Basin in 
the United States. Primary limiting factors appear to be temperature and spring flooding to elicit spawning. At the 
extreme northern limit of its range in Canada, Bigmouth Buffalo prefer a warm water habitat and may disperse further 
during a period of warmer chmate. It is tolerant of eutrophication and turbidity. 

Le buffalo a grande bouche, Ictiobus cyprinellus, est rare au Canada ou son aire de repartition est sporadique et 
extremement Hmitee. Dans une petite region de Saskatchewan, les populations sont suffisamment importantes pour 
justifier une exploitation commerciale de petite envergure, mais ailleurs, au Manitoba et en Ontario, elles sont 
extremement reduites et sporadiques si Ton en juge d'apres les tres rares specimens qui ont ete captures. Par contre, 
I'espece est repandue et abondante dans tout le bassin du Mississippi aux Etats-Unis. Les principaux facteurs limitants 
semblent etre la temperature et les crues printanieres qui conditionnent la fraye. A I'extreme nord de son aire de 
repartition au Canada, le buffalo a grande bouche prefere un habitat caracterise par une eau chaude, et pourrait se 
disperser davantage en periode de cHmat plus clement. II tolere I'utrophisation et la turbidite. 

Key Words: Bigmouth Buffalo, Ictiobus cyprinellus, buffalo a grand bouche, Catostomidae, buffalofishes, rare and 
endangered fishes, status, Canada. 



The Bigmouth Buffalo, Ictiobus cyprinellus 
(Valenciennes 1844), is characteristically a plains 
sucker (Figure 1) with an extremely limited and 
disjunct distribution in Canada. It typically 
inhabits slow, sluggish or still waters of larger 
rivers, oxbow and flood plain lakes, reservoirs, and 
lakes (Scott and Grossman 1979; Lee et al. 1980). A 
large deep-bodied, laterally compressed sucker, it 
closely resembles other buffalofish, {Ictiobus sp.), 
and is difficult to separate from Ictiobus niger, the 
Black Buffalo, the only other buffalofish found in 
Canada (Smith 1979). Comparative characters of 
three species of Ictiobus described by Crossman 
and Nepszy (1979) may assist identification. 
Ictiobus cyprinellus also resemble Carp (Cyprinus 
carpio) and members of the genus Carpiodes 
(Scott and Crossman 1973; Trautman 1981). 

Bigmouth Buffalo are described as robust with a 
slightly compressed body. The back is not highly 
elevated. The large oblique mouth and thin lips are 
key characters for identification. Usually, there are 
24 to 28 dorsal rays and nearly 100 gill rakers. The 
lower pharyngeal teeth are thin and weak, being 
more than twice as high as wide (Crossman and 
Nepszy 1979). Bigmouth Buffalo have a promi- 
nent, complete, almost straight lateral line with 
between 32 to 43 scales (Pfleiger 1975; Trautman 



1981); however, Canadian specimens have lateral 
line scale counts of 39 to 41 (Scott and Crossman 
1973). Fin ray counts are also highly variable with 
Canadian specimens having counts at the upper end 
of the range. 

The colouration is generally described as follows: 
back, upper side and head, dull brown to olive, sides 
lighter to yellowish, ventral surface white with dusky 
to gray fins. The colour may be light bluish-grey 
(Harlan and Speaker 1956; Trautman 1981). 
Reported differences in colour during spawning are 
described as follows: upper side more coppery, pale 
green to bluish, bluish; dorsal and caudal fins drab 
gray; the pelvic fins lighten and the pectoral fins 
become dull white (Harlan and Speaker 1956; Scott 
and Crossman 1973). Breeding males are reported to 
be slightly darker (Pfleiger 1975). 

Noticeable nuptial tubercles were reported for the 
first time by Morris and Burr (1987) from male 
Ictiobus cyprinellus in Illinois. Tubercles were 
distributed over much of the body including head, 
caudal peduncle and fin rays. Previously, nuptial 
tubercles had not been reported for either sex of this 
species (Scott and Crossman 1973; Pfleiger 1975). 

Potentially extremely large, a specimen from 
Spirit Lake, Iowa, was reported to weigh 36.3 kg (80 
lb) [Harlan and Speaker 1956]. Most adults, 



* Vulnerable status approved and assigned by COSEWIC 1 1 April 1989. 

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Vol. 104 




Figure \. Bigmouth Buffalo, Ictiobus cyprinellus [drawing by A. Odum from Scott and 
Grossman (1973) by permission]. 



however, are under 4.5 kg (10 lb) and less than 410 
mm (20 inches) in length (Smith 1979). In Canada, 
they are usually 254 to 457 mm (10 to 18 inches) in 
length (Scott and Grossman 1973). 

Distribution 

North America: The Bigmouth Buffalo is 
widespread throughout the Mississippi River Basin 
but the only drainage system of the Atlantic slope 
occupied is that of the Great Lakes system. 

It is found from Lake Erie south through the Ohio 
and Mississippi River basins (in Ohio, West 
Virginia, Kentucky, and Tennessee) to the 
Tennessee River in northern Alabama, south 
through Mississippi and near the Gulf of Mexico in 
Louisiana, northwest through eastern Texas and 
Oklahoma (rare), north through Kansas, Nebraska, 
South Dakota and North Dakota and west to the 
Milk River drainage in central Montana. From 
Illinois in the Mississippi River drainage, it ranges 
northwest through western Minnesota and north in 
the Red River into Manitoba and west into 
Saskatchewan (Figure 2) (Johnson 1963; Scott and 
Grossman 1973; Lee et al. 1980). Recent collections 
place it in the Lake of the Woods, northwestern 
Ontario (see Canadian Distribution below). It is 
absent from the Lake Superior drainage (Eddy and 
Underbill 1974). 

Introduced into Arizona, it is well established in 
some impoundments and it also occurs in the Gila 
River drainage. It has been introduced into 
California in the Los Angeles aqueduct system as 
well (Johnson 1963). 

The Bigmouth Buffalo appears to be native to 
Lake Eric. A specimen was reported from 
Rockport, Ohio by Kirkland as early as 1854 
(specimen in the Harvard Museum of Comparative 
Zoology). Hubbs (1930) thought it sporadically 
present in the Lake Eric watershed prior to 1900. 
C"urrcnt population levels arc likely a result of the 
United States (iovernment stocking of Bigmouth 
Buffalo in western Lake Erie and Sandusky Bay in 



approximately 1920. By 1949, numbers had 
increased to the level whereby Bigmouth Buffalo 
contributed substantially to the commercial fishery 
in the United States portion of Lake Erie (Trautman 
1981). 

Canada: In Canada, the distribution of Ictiobus 
cyprinellus is disjunct and limited. It occurs in: 
western Lake Erie, Lake St. Clair the St. Clair 
River and some tributaries, Lake Ontario, possibly 
Lake of the Woods in Ontario, the Red and 
Assiniboine Rivers and recently Delta Marsh 
(Lake Manitoba) in Manitoba, and west in the 
Qu'Appelle River system in central Saskatchewan 
(Figure 3). 

The first published record for this species in 
Canada may have been that of Gilchrist listed 
under the name Ictiobus bubalus (Scott and 
Grossman 1973). 

Ictiobus cyprinellus may have dispersed into 
Lake Erie postglacially or, more recently, because 
of changes in physical barriers and short-term 
climatic changes. It probably occupied a 
Mississippi refugium and may have used the Grand 
Valley outlet and glacial Lake Warren as an avenue 
of dispersal. In recent historical times, however 
there are many documented crossovers. Dispersal 
of the Bigmouth Buffalo is possibly temperature 
dependent as it may be restricted to the 22.5° C July 
isotherm (N.E. Mandrak, personal communica- 
tion). 

Although the species has been known from the 
Ohio waters of Lake Erie since the mid- 1 800s, it 
was not taken in the Canadian waters of Lake Erie 
until 1957 when an individual weighing 8.2 kg (18 
lb 2 07.) was caught by a commercial fisherman in 
Long Point Bay [Royal Ontario Museum (ROM) 
1971] (Scott 1957). It has been collected on only 
two subsequent occasions; from Port Dover in 
1968 [ROM 27749] and from Long Point Bay in 
1972 [ROM 28266). 

Why Bigmouth Buffalo should be well 
established in the United States' waters of Lake 



1990 



GOODCHILD: STATUS OF THE BiGMOUTH BUFFALO 



89 




Figure 2. North American distribution of the Bigmouth Buffalo, Ictiobus cyprinellus (adapted from Lee 
et al. 1980). 



Erie and apparently uncommon (sparse) in the 
Canadian part of the lake might be explained by 
the temperature requirements of this species. Its 
gradual movement into Canadian waters may be 
the result of an overall chmatic warming trend. 
Increased turbidity in the lake may also provide 
access to a tolerant species such as the Bigmouth 
Buffalo. In a recent study, fish species having the 
potential to invade the Great Lakes during a period 
of climatic warming were identified based on a 
discriminant function analysis of ecological 
characteristics. The Bigmouth Buffalo was one of 
the species that proved to have significant invasion 
potential (Mandrak 1989). 

Recently, the Bigmouth Buffalo has been 
collected from the St. Clair River by the Ontario 
Ministry of Natural Resources (OMNR) during 
electrofishing in "areas of concern" in the Great 
Lakes, as designated by the International Joint 
Commission. One specimen was electrofished 
from the northern part of the St. Clair River 
towards Lake Huron [A0CMNR86]. Two other 
specimens were collected at the outlet of the 
Lambton Generating Station [A0CMNR86]. 
Three additional specimens were collected from 
the Thames river at Jeanettes Creek [ROM 36582]. 

Its occurrence in the Bay of Quinte, Lake 
Ontario [ROM 37952] may be the resuh of an 
accidental introduction. The species is imported 
live from the United States and is often sold in 
Chinese fish markets. OMNR staff (Maple 



District) collected live specimens of several species 
not indigenous to Ontario, including Ictiobus 
cyprinellus, at Toronto markets (G. Goodchild, 
Fisheries Branch, Ontario Ministry of Natural 
Resources, Toronto, Ontario; personal communi- 
cation). A specimen purchased at a Toronto 
market is in the ROM collection [ROM 0]. 

Collections of Bigmouth Buffalo from the Lake 
of the Woods, in northwest Ontario represent a 
significant extension of the range of this species in 
Ontario. A specimen was collected 27.4 km north 
of Rainy River in June 1973 [NMC: Canadian 
Museum of Nature 70-0125A]; and another 
specimen was collected from Whitefish Bay in 
October 1973 [ROM 30431]. 

Bigmouth Buffalo were absent from collections 
in the southern part of Manitoba for many years. 
They were reported from the Red River of the 
North by Eigenmann (1895). Hinks (1943) 
reported only one specimen of 12.7 kg (28 lb) from 
southern Manitoba and Scott and Grossman 
(1973) considered it rare to absent in the Red and 
Assiniboine Rivers. In 1982 and 1983, however, 
Bigmouth Buffalo were collected in Delta Marsh at 
the south end of Lake Manitoba (Stewart et al. 
1985). This represent a significant extension of the 
range of this species in Manitoba and also verifies 
its continued existence in the Province. Recent 
collections from Lower Devil Lake in the Lake 
Winnipeg drainage [ROM 40369] and from the 
Red River near East Selkirk [ROM 35118] provide 
further evidence of a more extensive distribution. 



90 



The Canadian Field-Naturalist 



Vol. 104 




Figure 3. D'lstnhution oi Ictiobus cyprinellus in Camaia. 



The Assiniboine River Floodway which diverts 
floodwater from the river into Lake Manitoba near 
Delta Marsh was probably the avenue of dispersal 
used by a number of species (including Bigmouth 
Buffalo) recently reported from Lake Manitoba. 
The floodway was opened briefly in 1970 but the 
west dike, which allows excess water to flow 
directly into Delta Marsh, was not opened until 
1974 (Stewart et al. 1985). Further substantiation 
is the recent report of Ictiobus cyprinellus from the 
Assiniboine River in Manitoba (Grossman and 
McAllister 1986). 

Bigmouth Buffalo may have originally dispersed 
into Manitoba from the Mississippi River via the 
Red River. Stewart et al. (1985) suggest that it 
entered the Red River after 2000 BP. Glacial Lake 
Agassiz was probably not involved in the dispersal 
as Bigmouth Buffalo likely invaded much later 
than the recession of the ice flows due to limiting 
water temperature (N.E. Mandrak, personal 
communication). Lake Agassi/ is generally 
considered to have been formed earlier than other 
glacial waters (Radforth 1944) and Bigmouth 
Buffalo is a warm water species limited to the 
southern part of the Hudson Bay watershed. There 
are many species which occur in the Red River but 



not in the Souris River. Some have moved 
upstream into the English-Winnipeg system which 
possibly explains its occurrence in the Lake of the 
Woods. Others such as the Bigmouth Buffalo, have 
dispersed into the Assiniboine-Qu'Appelle system 
and become established (Grossman and McAllister 
1986). 

In Saskatchewan, Bigmouth Buffalo are 
abundant in the eight larger lakes of the 
Qu'Appelle drainage system (Johnson 1963; Atton 
1983). Extensive seining of the Frenchman, Souris, 
and Assinboine river drainages in Saskatchewan 
has failed to disclose its presence there. Despite its 
occurrence in the Milk River of Montana, part of 
the Missouri drainage, it apparently does not occur 
in the Missouri drainage of Ganada and is 
excluded from the distributional list of fishes in the 
Ganadian section of the Missouri drainage 
(Willcock 1969). 

Dispersal into Saskatchewan was probably 
through postglacial connections at the headwaters 
of the Mississippi and Red river systems. The 
headwaters of these systems are near Big Stone 
Lake and Lake Traverse on the South Dakota- 
Minnesota border and they were, apparently, 
connected during late glacial time. Today, the 



1990 



GOODCHILD: STATUS OF THE BiGMOUTH BUFFALO 



91 



divide may still be incomplete and headwaters may 
merge during spring floods (Johnson 1963). 

Protection 

Although it is considered to be of special 
concern in Manitoba (Johnson 1987), no specific 
protection for this species exists in Canada other 
than that afforded by habitat sections of the 
Fisheries Act. 

Bigmouth Buffalo are not considered a game 
species, and therefore, are not protected as such. 
The species is rarely taken on a hook and line and is 
not a generally sought after species in Canada. 
Thus, it is not hkely to be susceptible to angling 
pressure. 

While commercially important in the United 
States portion of Lake Erie, it has not contributed 
to the Canadian catch and is not specifically 
protected by Canadian regulations (J. Tilt, 
Fisheries Branch, Ontario Ministry of Natural 
Resources, Toronto, Ontario; personal 
communication). 

Population Sizes and Trends 

There has been insufficient assessment to 
accurately judge whether Canadian populations of 
Bigmouth Buffalo are increasing or decreasing in 
abundance. In the Qu'Appelle system, Saskatche- 
wan, they are abundant enough to support a small 
commercial fishery. Elsewhere in Canada, 
Bigmouth Buffalo exist in extremely low numbers. 
The species was thought to be absent from 
Manitoba for many years. Although recent 
collections from the Red River, Lower Devil Lake, 
and Delta Marsh {see Distribution above), indicate 
its continued existence in Manitoba, the numbers 
are too low to be encouraging. Populations in 
Manitoba may be extremely tenuous. 

Bigmouth Buffalo appear to be on the increase 
in southwestern Ontario. Its distribution may be 
expanding slightly based on recent collections 
from the Thames River and the St. Clair River but 
very few individuals have been collected and no 
population trends can be determined. 

In Saskatchewan, there is an erratic commercial 
fishery for Bigmouth Buffalo. The population in 
the Qu'Appelle drainage, Saskatchewan, is 
probably dependent on the sporadic production of 
strong year-classes (Johnson 1963). The 1948 year- 
class predominated in lakes studied during the mid 
1950s. Johnson (1963) found that it contributed 
88% of Bigmouth Buffalo caught in 1955 and 
approximately 82% of those caught in 1956. In 
fact, the 1948 year-class outnumbered all other 
year-classes in all the lakes sampled and no 
appreciable spawning or fry survival occurred in 
the years 1949 to 1954. Poor reproduction 
occurred in 1955 as well, but there was somewhat 



better success in 1956. Overabundance of the 1948 
year-class may have resulted in a much slower 
growth rate due to intra-specific competition. Late 
maturity of Bigmouth Buffalo was probably 
directly related and self-limiting, affecting 
reproductive rate in subsequent years. Dramatic 
changes in year-class strength may reflect a lack of 
specific requirements which may, in turn, make the 
species particularly vulnerable. 

Despite extensive trap netting in 1983, no 
juvenile Bigmouth Buffalo were caught in the 
Delta Marsh, Manitoba (Stewart et al. 1985). This 
further supports the theory that there is an extreme 
variation in year-class abundance for this species, 
at least in the northern extremity of its range. 

Large fluctuations in year-class strength have 
been noted in other studies of Bigmouth Buffalo 
populations. In North Dakota, dramatic increases 
in previously strong year-classes were attributed to 
loss of optimal spawning areas (Willis 1978). In 
Lake Oahe, a Missouri River reservoir, Bigmouth 
Buffalo were dominated by three strong year- 
classes. The 1962 year-class was exceptional, 14 to 
16 times more abundant than the 1965 year-class at 
ages IV and V respectively (Moen 1974). 
Recruitment appears to be sporadic among 
buffalofish populations in other waters as well. 

In the United States, Bigmouth Buffalo 
populations tend to be relatively stable and there 
are few reports of drastic declines. Generally, the 
species appears to be maintaining or increasing in 
population numbers and distribution. Its decrease 
in abundnace and distribution in Minnesota and 
North Dakota is an exception (Eddy and Underbill 
1974). 

If size, as well as number of individuals, is 
considered, the species is sometimes incredibly 
abundant. Along the Mississippi River and its 
tributaries there is a significant inland fishery, and 
it is of considerable commercial importance in 
many states (Scott and Crossman 1973; Becker 
1983). Although abundance in Minnesota has 
decreased, tremendous numbers continue to be 
taken by commercial fishermen in Lac La Croix 
during summer and from the Mississippi River 
near Winona in winter. In the early 1900s to mid- 
1950s, spawning runs in central and southern 
Minnesota contained similar vast numbers (Eddy 
and Underbill 1974). 

Habitat 

Bigmouth Buffalo may inhabit deeper pools of 
large streams, shallow overflow ponds, natural 
lowland lakes and man-made impoundments; 
usually in moderate to slow current (Pfleiger 1974; 
Trautman 1981). Pfleiger (1975) perceived that 
they occur in schools at midwater or near the 
bottom. In Saskatchewan, Bigmouth Buffalo 



92 



The Canadian Field-Naturalist 



Vol. 104 



occupy shallow lakes with a maximum depth 
varying from 5 m to 30 m. Capture data indicate 
they prefer water shallower than 5 m and the 
maximum depth of capture was 8.5 m (Johnson 
1963). They are common in Lake Poinsett, South 
Dakota, which has a maximum depth of 5.5 m 
(Starostka and Applegate 1970). In Wisconsin, 
they are encountered most frequently in water 
more than 1.5 m deep over substrates of mud, silt, 
sand, gravel, clay and rubble (Becker 1983). 

The species is noteworthy for its tolerance to 
turbidity which is evidently higher than in other 
species of buffalofishes (Pfleiger 1975). Its sustained 
abundance in the Ohio River since 1925 suggests 
that species with large terminal mouths may be more 
suited to turbid water conditions and rapid silting of 
stream bottoms than species with smaller, inferior, 
mouths such as Smallmouth Buffalo (Trautman 
1981). In Lake Erie, Bigmouth Buffalo are most 
abundant in the upper half of Sandusky Bay where 
the water is often extremely turbid. 

The species exhibits a preference for warm, 
highly eutrophic, waters. Bigmouth Buffalo are 
abundant in Echo and Pasqua Lakes in Saskatche- 
wan, which are described as highly eutrophic with 
heavy standing crops of benthic fauna and 
plankton. Dense blue-green algal blooms occur 
frequently and thermal conditions range from 
almost complete mixing throughout summer to the 
development of a pronounced thermocline 
(Johnson 1963). In Clear Lake, Iowa (a turbid 
prairie lake), Bigmouth Buffalo were most 
abundant in vegetated areas (Stang and Hubert 
1984). Reservoirs in Arizona stocked with 
Bigmouth Buffalo in the 1920s, are described as 
basically clear, except near areas of silt or high 
nutrient input where planktonic algae may 
produce high seasonal turbidity. In Roosevelt 
Lake and Apache Lake, Arizona, freezing rarely 
occurs and surface temperatures may exceed 30° C 
with pronounced summer stratification. These 
lakes are also somewhat saline, indicating that 
Bigmouth Buffalo are tolerant of a mild degree of 
salinity (Minckley et al. 1970). Roosevelt Lake has 
fluctuating water levels and conditions of 
turbidity. Bigmouth Buffalo are abundant in Lake 
Poinsett, the largest natural lake in South Dakota. 
It is eutrophic and devoid of emergent aquatic 
vegetation (Starostka and Applegate 1970). 

Bigmouth Buffalo arc able to endure low oxygen 
tensions and high water temperatures. In a test, 
oxygen was reduced to 0.9 mg.L' and no deaths 
occurred (Gould and Irwin 1962). The maximum 
temperature recorded during capture of this 
species is 3 1 .7'^'C (89'^ F) (Proffitt and Bcnda 1971). 

Spawning occurs in shallow bays or small 
tributary streams (Eddy and Underbill 1974). 
Bigmouth Buffalo from Lake Erie have been 



observed spawning in tributary streams of 
Sundusky Bay, even ascending small ditches. 
Individuals exceeding 1 1 .3 kg (25 lb) were captured 
in shallow ditches only a few feet wide (Trautman 
1981). They readily invade marshes, tributary 
streams, ditches, and backwaters during spring 
flooding (Johnson 1963). 

Bigmouth Buffalo share an environment with 
predacious fishes such as Northern Pike {Esox 
lucius). Black Bullhead {Ictalurus melas). Burbot 
(Lota lota). Yellow Perch {Perca flavescens), and 
Walleye {Stizostedion sp). In Pasqua Lake, 
Saskatchewan, the species most often found 
associated with Bigmouth Buffalo were Yellow 
Perch, White Suckers {Catostomus catostomus) 
and Spottail Shiners {Notropis hudsonius) 
[Johnson 1963]. 

Distribution of Ictiobus cyprinellus in Canada is 
restricted and localized. They are apparently 
tolerant of changes in habitat associated with 
turbidity and eutrophication but the amount of 
critical habitat has not been estimated. Habitat 
change, resulting from human interference, 
possibly increases the amount of suitable habitat 
for Bigmouth Buffalo. For instance, it fares well in 
man-made water bodies such as reservoirs. 
Protection of areas of suitable habitat does not 
appear to be necessary at this time. 

General Biology 

Reproductive Capability: Johnson (1963) 
provides considerable information on the life 
history of the Bigmouth Buffalo in Canada. It 
appears, however, that the individuals he studied, 
from lakes in Saskatchewan, matured later and 
had slower rates of growth than in most other 
areas. Bigmouth Buffalo may have considerable 
variability in age of maturation. Harlan and 
Speaker (1956) report sexual maturity is attained 
in the third year and found that pond-reared fish 
spawned at four years of age. Although age of 
spawning has been reported from as young as one 
year of age (Johnson 1963, after Swingle 1957), in 
Pasqua Lake, Saskatchewan, maturity was 
delayed. Gonads of 360 females less than 482 mm 
(19 inches) long, probably 7 to 9 years of age, were 
examined and only 19 were mature. The largest 
immature fish seen was a female of 475 mm (18.7 
inches). This is probably the corollary of slow 
growth due to an extremely dense year-class (1948) 
and the cooler temperatures at this latitude. Males 
reach sexual maturity at a considerably smaller 
size than females. They begin to mature after 
attaining a length of 305 mm ( 12 inches) and most 
males arc mature in the range of 356 to 379 mm 
(14.0 to 14.9 inches) [Johnson 1963]. 

Burr and Heidingcr (1983) observed groups of 
three or more individuals spawning in 0.5 to 0.75 m 



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GOODCHILD: STATUS OF THE BiGMOUTH BUFFALO 



93 



deep water in Crab Orchard Lake, Illinois. 
Usually, two tuberculate males aligned alongside 
one female in a "rush" along the water surface. 
Spawners then sank to the bottom and sometime 
assumed a vertical posture. Adhesive eggs were 
scattered over decaying vegetation and left 
unattended. Eggs are reportedly scattered and 
abandoned in shallow water over plant-debris, to 
which the eggs adhere until they hatch (Cross 1967; 
Eddy and Underbill 1974). In the Missouri River, 
however, fishes thought to be Bigmouth Buffalo 
were observed to be spawning by broadcasting 
eggs over rocks in water so shallow that the backs 
of the spawning fish were exposed (Pfleiger 1975). 

Johnson (1963) found a storng relationship 
between spring flooding and reproductive success 
in spawning Bigmouth Buffalo. The strongest 
year-class was produced in 1948 when the greatest 
May run-off occurred. Earlier studies, involving 
pond culture of Bigmouth Buffalo found that the 
introduction of fresh water was necessary to 
activate spawning. If heavy run-off is the major 
factor governing reproductive success, then one 
would expect population peaks associated with 
flood years. 

Bigmouth Buffalo spawn in spring during April, 
May, and as late as June in the northern part of the 
range in Saskatchewan. The sex ratio during 
spawning of large fish was two females to three 
males but, later in the season, was one female to 
five males (Johnson 1963). Optimal water 
temperatures for spawning are between 15.6° to 
18.3° C (60° to 65° F), but eggs are laid in water up 
to 26.7°C (80° F) [Johnson 1963]. In Ilhnois, 
spawning occurred at the end of April in water 8° 
to 10°C (Burr and Heidinger 1983). Eggs hatch in 
about ten days at 16.7°C (62° F) [Eddy and 
Underbill 1974]. Johnson (1963) estimated the 
number of eggs contained in a 665 mm (26.2 inch) 
long individual at approximately 750 000. Over 
400 000 eggs were estimated to be contained by a 
4.5 kg (9.9 lb) female (Harland and Speaker 1956). 
Size of mature eggs range from 1.2 to 1.8 mm in 
diameter (Scott and Grossman 1973). 

Length of young-of-the year from Ohio range 
from 43 to 102 mm (1 .7 to 4.0 inches) in September 
and individuals reach 127 to 178 mm (6.0 to 7.0 
inches) by about one year (Trautman 1981). 
Bigmouth Buffalo from Minnesota and Tennessee 
were compared up to age 7 and those from 
Minnesota were consistently longer at each age 
group (Carlander 1950). 

Ictiobus cyprinellus from the Qu'Appelle River 
grew more rapidly than those from lakes in 
Saskatchewan, attaining an average length of 71 
mm (2.8 inches) in the first summer. In Pasqua 
Lake, Saskatchwan, the largest young-of-the-year 
captured was 63 mm (2.5 inches) long in August 



1956. The unique population Johnson studied in 
Pasqua Lake was severely affected by the 
extremely numerous 1948 year-class which, 
combined with the northern climate, resulted in a 
stunted population. Growth was exceedingly slow 
compared with that in Reelfoot Lake, Tennessee, 
where 3-year-old fish are nearly as long as a 9-year- 
old Pasqua Lake fish. Dominance of three strong 
year-classes (1959, 1960, 1962) apparently resulted 
in a reduced growth rate of Lake Oahe Bigmouth 
Buffalo, as well. Individuals of the 1962 year-class 
averaged 10 to 11% shorter and 36% lighter than 
those of the 1959 year-class (Moen 1974). 

Adults are usually 590 to 1180 mm (15.0 to 30 
inches) long and weigh from 0.9 to 13.6 kg (2 to 30 
lb), ahhough lengths of 1370 to 1570 mm (35 to 40 
inches) and weights of 18.1 to 27.2 kg (40 to 60 lb) 
have been reported in Ohio. Maximum reported 
weight is 36.3 kg (80 lb) [Trautman 1981]. 

Johnson (1963) was confident in assigning age 
groups for fish up to 10 years of age. Age 
determinations for older fish were progressively 
less reUable with increasing age. Moen (1974) 
found that age determinations became increasingly 
difficult beyond age V, especially for males. 
Bigmouth Buffalo from lakes in Saskatchewan 
included individuals estimated to be up to 20 years 
of age [696 mm (27.4 inches)]. Bigmouth Buffalo 
may have very long life spans judging from the 
extremely large size reportedly attained by some 
individuals. 

Species Movement: The Bigmouth Buffalo is 
not generally considered to be a migratory species. 
It does, nonetheless, migrate in order to spawn in 
flooded marshes and tributary streams and may 
travel long distances in order to find suitable sites 
(Eddy and Underbill 1974; Cooper 1983). 
Although Johnson (1963) did not investigate 
movement, he did allude to spawning migration by 
indicating the species had been captured during 
spring in locations presumably associated with 
breeding movements. 

Backer (1983) suggests that, in addition to 
migration in upstream spawning runs in spring, 
Bigmouth Buffalo may also participate in runs in 
September and October when high flood waters 
are present. In South Dakota, movement of 
marked fish was extensive. The maximum distance 
traveled was 379 km and the maximum rate of 
travel was 6.5 km per day (Moen 1974). 

Behaviour I Adaptability: Starostka and Apple- 
gate (1970) determined that Bigmouth Buffalo 
greater than 236 mm exhibit little, if any, food 
selectivity and described it as an indiscriminant 
planktivore. In most instances, they found that 
subadults and adults consumed plankton in the 
same proportion as found in the limnetic 



94 



The Canadian Field-Naturalist 



Vol. 104 



environment, ingesting primarily Daphnia and 
other cladocerans, copepods, and blue-green 
algae. Analysis of stomach contents indicated a 
succession in food habits as fry had consumed 
primarily benthic organisms. 

The enlarged head and mouth, and structure of 
the gill rakers are indicative of planktonic feeding 
(Minckley et al. 1970). Stomach contents, from 
early studies, supported the conclusion that the 
Bigmouth Buffalo were predominantly plankton 
feeders. However, studies in Arizona indicate a 
near bottom feeding habit (Johnson and Minckley 
1972); analysis of the diet of young, juvenile, and 
adult Bigmouth Buffalo in Pasqua and Echo 
Lakes, Saskatchewan, also revealed that many 
animals classed as bottom fauna are often taken as 
food (Johnson 1963). The organisms include 
chironomid larvae, miscellaneous insects, mol- 
luscs, amphipods and ostracods. The largest 
volume of food, however, was comprised of 
cladocerans and copepods, normally thought of as 
planktonic, yet both groups have members classed 
as creepers which spend a great deal of their life on 
a substrate and Johnson (1963) demonstrated 
these contributed to the diet of Bigmouth Buffalo. 
He found a tendency to increased utilization of 
benthic invertebrates by larger fish, contrary to the 
results obtained by Starostka and Applegate 
(1970). Diet is probably influenced by availabihty 
of foods rather than by active selection. Bigmouth 
Buffalo occupy a food niche overlapping both 
benthic and planktonic feeding (Johnson 1963). 
Minckley et al. (1970) suggest that Bigmouth 
Buffalo may utilize the organisms churned up by 
their "bouncing" feeding movements. They have 
been observed in hatcheries swimming at an angle 
of 55° to the bottom "skipping" back and forth as 
they sucked up food pellets. Similar feeding 
behaviour was described for non-captive individu- 
als (Johnson 1963). 

Changes in the ecology of a water body, brought 
about by human disturbance, often result in a 
drastic alteration of the biota. As Bigmouth 
Buffalo appear to be opportunistic feeders, they 
would be expected to be able to utilize an altered 
fauna. In some Arizona lakes that contain non- 
native species of plankton and benthic organisms, 
Bigmouth Buffalo are well established. They are 
also relatively tolerant of increased turbidity and 
cutrophication which often accompanies anthro- 
pogenic changes (Minckley et al. 1970). 

Parasitic infections, particularly those which 
interfere with the feeding mechanisms of Bigmouth 
Buffalo may have detrimental effects on popula- 
tions. Johnson (1963) found young fish heavily 
infected with spores of Myxosporida enclosed in 
prominent white cysts on the gills. The smallest fish 
were emaciated. Hctoparasitic copepods (Argulus 



apendiculosus) were also found on individuals in 
Saskatchewan. These are the only parasites 
recorded for this species in Canada (Margolis and 
Arthur 1979). Hoffman (1967) listed the following 
parasites of Bigmouth Buffalo: Trematoda, 
Cestoda, Nematoda, Acanthocephala, leeches, 
Crustacea. Bigmouth Buffalo kept in holding 
tanks appeared to be particularly susceptible to 
parasitic infections (Becker 1983). 

There is a pronounced tendency to school during 
the summer. On warm still days, Bigmouth Buffalo 
spread out over the entire surface of lakes where 
they rest quietly in the upper 0.6 m (2 ft) of water. 
Often the dorsal fin projects above the surface. The 
behaviour is never observed on windy days. 
Commercial fishermen take advantage of this 
congregating behaviour by setting barrier nets 
(Johnson 1963). 

The systematics of Ictiobus cyprinellus have not 
been studied in detail. There is no significant 
sexual dimorphism (Phillips and Underbill 1971). 
Species of Ictiobus appear to be a closely related 
group. In a biochemical study of the phylogenetic 
relationships of catostomid fishes, results 
indicated that there is close affinity between 
Ictiobus bubalus and Ictiobus cyprinellus and they 
hybridize naturally (Johnson and Minckley 1972; 
Bussjaeger and Briggs 1978). Ictiobus cyprinellus 
and Ictiobus niger have been artificially hybridized 
and progeny cultured (Lee et al. 1980, after 
Stevenson 1964). 

Limiting Factors 

Many factors found to be limiting to popula- 
tions of other fishes apparently have little effect on 
Bigmouth Buffalo. Although they may share an 
environment with many large predacious fishes, 
there is little evidence of predation on young 
Bigmouth Buffalo, and certainly, large adults are 
probably free from any predation (Johnson 1963; 
Scott and Crossman 1973). The gibbous shape, 
combined with large size, probably makes them 
difficult to swallow. 

Turbidity, siltation and cutrophication cannot 
be tolerated by many species of fish, but the 
Bigmouth Buffalo is evidently able to thrive under 
these conditions. It prefers warm, slow flowing, 
shallow waters with abundant bottom fauna and 
plankton. Ecological changes resulting from 
human activities tend to produce aquatic habitats 
of this nature and thus may even increase the 
amount of suitable habitat for Bigmouth Buffalo. 
Certainly, they thrive in impoundments and, in 
some cases, have become important in the 
commercial harvest of these reservoirs. 

Prime among the limitations on this species is 
the requirement for spring flooding to elicit 
spawning. The very successful 1948 year-class 



1990 



GOODCHILD: STATUS OF THE BiGMOUTH BUFFALO 



95 



which comprised most of the individuals studied 
by Johnson (1963) was produced in a year when 
there was significant spring run-off. Little 
successful reproduction was observed in the years 
when freshets did not occur. Therefore, channeli- 
zation and flood control measures may have 
detrimental effects on reproductive success. 

Nevertheless, an extremely successful reproduc- 
tive rate may be self-limiting. Overabundance of 
the species resulting in a high level of intraspecific 
competition may cause stunting and late maturity 
of individuals. Subsequent year-class strength may 
be extremely low or non-existent. In his studies of 
lake populations in Saskatchewan, Johnson (1963) 
surmised that the uniquely successful 1948 year- 
class was partly responsible for a distorted growth 
rate and misleading appraisal of size and age of 
sexual maturation. Strong and weak year-classes 
are very apparent (Scott and Grossman 1973) and 
females in Saskatchewan may not spawn every 
year. 

The degree of interspecific competition has not 
been studied. There are indications though that the 
Bigmouth Buffalo may have a selective advantage 
in occuying a food niche that overlaps both benthic 
and limnetic feeding. 

Heavy parasitic infestations, particularly by 
Myxosporida which are enclosed in cysts on the 
gills, may severely debilitate populations, due to 
interference with feeding mechanisms. The gill 
rakers of Bigmouth Buffalo are specialized for 
plankton feeding (Johnson 1963; Starostka and 
Applegate 1970). 

Since Bigmouth Buffalo are generally consi- 
dered to be a "rough fish", they are not protected 
by regulations that protect other more favoured 
species. Populations have been indiscriminately 
eliminated during attempts to remove unwanted 
species such as carp. Toxicants were used in 
Wisconsin which eradicated all fish species from 
the area treated. As a result, Bigmouth Buffalo are 
no longer present in many areas of the Upper Rock 
River drainage. Becker (1983) suggests that the 
classification of Bigmouth Buffalo in Wisconsin 
should be changed to sport fish status. 

Special SigniHcance of the Species 

The blood of Ictiobus cyprinellus and Ictiobus 
bubalus, in common with only a few other species, 
contains unique cells called secretory granulocytes 
(Chlebeck and Phillips 1969). The function of these 
cells is unknown. 

Bigmouth Buffalo are not usually considered 
sport fish and the species seldom takes a hook. In 
the lower Mississippi Valley, it is frequently taken 
on set lines baited with balls of dough (Jordan and 
Evermann 1923). In Wisconsin, fishermen seek the 
Bigmouth Buffalo with square dipnets from river 



banks and bridges or they pursue it on flooded 
marshes from rowboats and skiffs using spears 
(Becker 1983). Bigmouth Buffalo is becoming 
increasingly popular as a table fish and is an ethnic 
favourite. The meat is nutritious and considered to 
be superior to carp in taste. It is excellent when 
smoked (Becker 1983). 

Under managed conditions, buffalo fishes have 
a great potential for producing high-level protein 
and culture could be profitable. A single trial of 
Bigmouth Buffalo in a 1/25 ha pond resulted in a 
yield of 287 kg.ha' without fertilization of the 
pond or feeding of the fish (Cross 1967). Bigmouth 
Buffalo have been cultured in ponds since the early 
1900s. The culture of trout and buffalofish 
{Ictiobus sp.) expanded in the 1950s but culture of 
Ictiobus dwindled by the early 1960s. In Arkansas, 
in 1979, 4000 metric tonnes of buffalofish were 
marketed, however, only about 600 metric tonnes 
were pond-raised (Shelton and Smitherman 1984). 
It is currently a prized cultured species in eastern 
Europe and the USSR. Becker (1983) suggests that 
the use of ponds receiving treated effluent water 
from sewage plants and cooling ponds at power 
generation sites, should be considered for the 
culture of Bigmouth Buffalo. 

There is a significant commercial fishery for this 
species in the United States. Commercial 
fishermen from Iowa, Missouri, and Nebraska 
reported taking 120 327 kg of Smallmouth and 
Bigmouth Buffalo in 1976 and 1977. The species 
contributes a major portion of the commercial 
catch of the Mississippi River with yields of 672 
kg.ha' (600 lb/ acre surface area) not uncommon 
in some areas (Harlan and Speaker 1956). In just 
the Wisconsin portion of the Mississippi River, 
from 1960-1964, the total harvest of buffalos 
(combined species) was 1 257 000 kg. Production 
in the Wisconsin waters of the Mississippi River 
has continued to be substantial. In 1976, for 
instance, 243 962 kg were captured with a cash 
value of $115 634. Bigmouth Buffalo contribute 
the greatest portion of the catch. 

Ictiobus cyprinellus is considered to be of special 
concern in Manitoba but not in any states of the 
United States (Johnson 1987). Until recently 
Ictiobus cyprinellus was the only member of the 
genus Ictiobus to occur in Canada. In 1978, the 
Black Buffalo, Ictiobus niger, was first described, 
from the western end of Lake Erie in Canada 
(Crossman and Nepszy 1979). It has since been 
collected from several other locations in southwest- 
ern Ontario. It has also been considered by 
COSEWIC and has also been assigned a vulnerable 
status in Canada (Houston 1990). The Black Buffalo 
is considered to be of special concern in Kentucky, 
Mississippi, South Dakota, and West Virginia in the 
United States and is protected in Wisconsin (John- 



96 



The Canadian Field-Naturalist 



Vol. 104 



son 1987). So few specimens of Ictiobus other than 
Ictiobus cyprinellus are seen from the Great Lakes 
that the characteristic features of other large 
Ictiobus are poorly known and, since they have a 
pronounced tendency to hybridize, the species may 
be readily confused. 

Evaluation 

The Bigmouth Buffalo is apparently flourishing 
in the Mississippi Valley watershed. It is also 
particularly tolerant of environmental degradation. 
In the United States part of its range, it is abundant 
and not in jeopardy. 

The position of most Canadian populations, 
however, is extremely tenuous. Although Bigmouth 
Buffalo are indigenous, they are, apparently, at the 
extreme northern limit of their range in Canada. 
Except for a well established population in Sask- 
atchewan, the species exists in extremely low 
numbers elsewhere. Temperature may be the crucial 
limiting factor that restricts its distribution in 
Canada as there seems to have been ample opport- 
unity for Bigmouth Buffalo to disperse into 
Canadian waters, especially Lake Erie. There is some 
indication that Bigmouth Buffalo may disperse 
further into Canadian waters during a period of 
warmer climate. Its competitiveness may be en- 
hanced by certain types environmental degradation. 

Since the species has potential economic value, is 
at the fringe of its range, and generally occurs in low 
numbers, it should be classified as vulnerable (rare) 
in Canada. 

Acknowledgments 

The assistance of D. E. McAllister of the National 
Museum of Natural Science (now Canadian 
Museum of Nature), E. Holm, R. W. Winterbot- 
tom, and E. J. Crossman of the Royal Ontario 
Museum, G. E. Gale and G. A. Goodchild of the 
Ontario Ministry of Natural Resources, in 
providing access to records and reports was 
invaluable. 

The author is particularly grateful to N. E. 
Mandrak, University of Toronto, for his insight into 
the zoogeographic aspects of the distribution of this 
species. 

W. B. Scott, Huntsman Marine Laboratory, and 
E. J. Crossman, Royal Ontario Museum, gave 
permission to use the line drawing of Ictiobus 
cyprinellus from the text Freshwater Fishes of 
Canada, Bulletin 184, Fisheries Research Board, 
Ottawa, Ontario. 

Sincere thanks to R. R. Campbell, Department 
of Fisheries and Oceans, Canada, for his support 
and understanding in the preparation of this report. 

Preparation of the report was supported through 
financial assistance of the World Wildlife Fund 
Canada. 



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Scott, W. B., and E. J. Grossman. 1973. Freshwater 
fishes of Canada. Fisheries Research Board of Canada 
Bulletin 184. 966 pages. 

Shelton, W. L., and R. O. Smitherman. 1984. Exotic 
fishes in warmwater aquaculture. Pages 262-287 in 
Distribution, biology, and management of exotic fishes. 
Edited by W. R. Courtnay, Jr., and J. R. Stauffer. 
Johns Hopkins University Press, Baltimore, Maryland. 
430 pages. 

Smith, P. W. 1979. The fishes of Illinois. University of 
Illinois Press. 314 pages. 

Smith- Vaniz, W. F. 1968. Freshwater fishes of Alabama. 
Auburn University, Agricultural Experiment Station. 
21 1 pages. 

Stang, D. L., and W. A. Hubert. 1984. Spatial separa- 
tion of fishes captured in passive gear in a turbid prairie 
lake. Environmental Biology of Fish 1 1(4): 309-314. 

Starostka, V. J., and R. L. Applegate. 1970. Food 
selectivity of Bigmouth Buffalo, Ictiobus cyprinellus, in 
Lake Poinsett, South Dakota. Transactions of the 
American Fisheries Society 99(3): 571-576. 

Stewart, K. W., \. M. Suthers, and K. Leaves- 
ley. 1985. New fish distribution records in Manitoba 
and the role of a man-made interconnection between 
two drainages as an avenue of dispersal. Canadian 
Field-Naturalist 99(3): 317-326. 

Trautman, M. B. 1981. The fishes of Ohio with 
illustrated keys. Revised edition. Ohio State University 
Press, Columbus, Ohio. 782 pages. 

Willis, D. W. 1978. Investigations of population 
structure and relative abundance of year-classes of 
buffalo fishes, Ictiobus spp. in Lake Sakakawea, North 
Dakota. M.Sc. thesis, North Dakota University, Grand 
Forks, North Dakota. 

Accepted 10 October 1989 



Status of the Black Buffalo, Ictiobus niger, in Canada* 

J. Houston 

374 Fireside Drive, Woodlawn, Ontario KOA 3M0 

Houston, J. 1990. Status of the Black Buffalo, /cr/ofew^^/ger, in Canada. Canadian Field-Naturalist 104(1): 98-102. 

The Black Buffalo, Ictiobus niger, has only recently been reported from Canadian waters in the western end of Lake 
Erie. One earlier United States record for western Lake Erie is also known. The species is less common than the other 
buffalofishes and appears to have declined or been extirpated in portions of its United States range, but has extended 
its range elsewhere through introductions. The lower Great Lakes, however, appear to be within the native range. The 
species is rare in Canadian waters. 

Ce n'est que recemment que le buffalo noir, Ictiobus niger, a ete signale dans les eaux canadiennes a I'extremite ouest 
du lac Erie. On connait egalement une mention americaine anterieure pour Fouest du lac Erie. L'espece est moins 
repandue que les autres buffalos et semble avoir diminue ou avoir ete deracinee dans certaines zones de son aire de 
dispersion aux Etats-Unis, mais elle a etendu son aire ailleurs grace a des introductions. Cependant, les lacs inferieurs 
des Grands Lacs semblent etre dans les limites de I'aire de dispersion naturelle. L'espece est rare dans les eaux 
canadiennes. 

Key Words: Black Buffalo, Ictiobus niger, buffalo noir, Catostomidae, suckers, buffalofish, rare and endangered 

fishes. 



The buffalofishes (genus Ictiobus) are large 
suckers (family Catostomidae) characteristic of the 
rivers, lakes and larger streams of the Mississippi 
drainage basin. They are easily separated from 
other genera of the family (Catostomidae are 
represented in North America by 10 genera and 
some 65 species, seven of the genera have Canadian 
distributions) by their large size and the shape of 
the head and body which more closely resemble 
those of Carp {Cyprinius carpio). They are easily 
distinguislb.d from the latter by lack of barbels, 
typical suckerlike mouth, and difference in colour. 
Amongst the suckers, buffalofishes most closely 
resemble the carpsuckers (Genus Carpiodes) from 
which large specimens can be distinguished by the 
thicker body and upward curve of the snout {see 
Clay 1962; Cross 1967), but juveniles may be 
confused with those of Carpiodes sp. 

The Black Buffalo, Ictiobus niger (Rafinesque 
1820), is the smallest of the buffalos, commonly 90 
to 60 cm in length and 0.5 to 4.5 kg in weight 
(Figure 1). The species can be distinguished from 
the other buffalos by the darker colour, thicker 
body and ventral mouth (Clay 1962). The dorsal 
surface is usually slate-grey to black, the sides 
brownish and the belly yellow to white. 

Distribution 

In Canada, the species was first described from a 
single specimen from the western end of Lake Erie 
[Crossman and Nepszy 1979; ROM (Royal 
Ontario Museum) 34562]. The Black Buffalo has 



also been reported from Boston Creek [Haldiman 
Norfolk County; 49°59'42"N, 80°16'18"W: 
OMNR (Ontario Ministry of Natural Resources) 
575 #42] of the Lake Erie drainage and in central 
Lake Erie [ROM 5397 1]. The OMNR has recorded 
collections from Carp Creek, a tributary of the 
Saugeen River in Grey County [44°08'36"N, 
80°32'00"W; 44° 09'36"N, 30° 54'24"W: OMNR 
#'s 83, 43, and 44 respectively]. There is one record 
of the species from the Niagara River near St. 
Catharines [43°19'27"N, 79°03'01"W: AOCMNR 
86: OMNR]. (Figure 3). 

These records have been checked by staff of the 
Royal Ontario Museum and the Boston Creek and 
Carp Creek records are catalogued as Castostomus 
sp. unidentified. The Niagara river specimen was 
returned to the water alive and cannot be verified 
(E. J. Crossman, ROM, Toronto, Ontario; 
personal communication). 

In the United States, the Black Buffalo is known 
from most large rivers and many smaller rivers of 
the Mississippi, Missouri and Ohio river basins 
(Figure 2). It has also been reported as rare and 
extirpated or depleted in the Calcasieu, Sabine, 
Bryos and Rio Grande drainages by the U.S. states 
concerned, but these may be introduced popula- 
tions or misidentifications of the more locally 
common Smallmouth Buffalo, Ictiobus bubalus 
(Hubbs and Lagler 1958; Shute 1980). 

In the Great Lakes, the species has only been 
reported from southern Lake Michigan and Lake 
Erie (Hubbs and Lagler 1958). Although 



*VulnL-tablc status approved and assigned by COSI WIC I I April I9K9. 



98 



1990 



HOUSTON: Status of the Black Buffalo 



99 




h 



Figure I. Black Buffalo, Ictiobus niger (drawn from photograph in Cross 1967). 



Trautman (1957) indicates that the lower Great 
Lake populations resulted from introductions, 
Hubbs and Lagler ( 1 998) disagree and consider the 
species to be native in these waters. Trautman 
(1957) does note the lack of early information on 
the species due to confusion with the Smallmouth 
Buffalo and its less common occurrence. He also 
indicates that hybrids between the species had been 
taken in Sandusky Bay, thereby supporting the 
existence of the species in Lake Erie. Moore (1968) 
hsted the species for Lake Erie. The recent 
Canadian collections (1975 to 1987) confirm the 
presence of the species in Lake Erie. 

Protection 

There is no specific protection for this species in 
Canada although general protection is available 
through the Fisheries Act. In the United States, the 
Black Buffalo is listed as a species of special 
concern in Kentucky, Mississippi, South Dakota 
and West Virginia. It has been Usted as protected in 
Wisconsin (Johnson 1987). 

Population Sizes and Trends 

Although Trautman (1957) indicated that the 
species was probably introduced to the lower Great 
Lakes, Hubbs and Lagler (1958) state they are 
native {see Distribution). Since the species is 
abundant in Ohio and can easily be confused with 
the Smallmouth Buffalo, especially smaller 
individuals (Smith 1979), it has probably gone 
unnoticed. Moreover, in the past, many fishermen 
and fisheries biologists believed this species to be a 
hybrid between the other two species and it was 
referred to as the Mongrel Buffalo Fish. The 
species is of no commercial interest and no attempt 
to ascertain populations in the Great Lakes have 
been made. Thus, it is probably native to Lake Erie 



and may have gone virtually undiscovered there 
because of its rarity, unimportance and similarity 
to the other species. 

The origins of this species in Canada and its 
occurrence in the Lake Huron drainage may be 
questionable, but based on its apparent rarity and 
other factors as discussed above, it is not 
unreasonable to assume that this fish may be native 
to the Lake Huron drainage as well and have 
previously gone unnoticed. On the other hand, it 
may have been introduced through release of bait 
fish. More information on the distribution of the 
species in Canada is required before this question 
can be resolved. However, the species appears to be 
native to Canadian waters, at least in Lake Erie, 
where it is relatively rare. 

In the United States, the species is still listed as 
common in some parts of the range, but has been 
depleted or extirpated in other parts (Smith 1979; 
Shute 1980). The distribution is sporadic, related 
to availability of suitable habitat and the species 
seems intolerant of pollution. On the other hand, 
they do adapt to impoundment conditions (Cross 
1967). Nowhere are they as abundant as the other 
two buffalos. 

Habitat 

No specific information is available on the 
habitat preferences of the species, but Black 
Buffalo are often found in association with the 
Smallmouth and Bigmouth {Ictiobus cyprinellus) 
buffalos. However, Trautman (1957) indicated the 
preferred habitat was intermediate to that of the 
other two. The Bigmouth Buffalo is often found in 
shallow turbid pools, overflow ponds and lowland 
lakes and has a high tolerance for turbid waters 
(Trautman 1957). The Smallmouth Buffalo, on the 
other hand, frequents less turbid waters and its 



100 



The Canadian Field-Naturalist 



Vol. 104 




45° 



25°_ 



i 
® 



Line encloses native distribution 
(Transplanted populations not mapped) 



105° 80° 

Figure 2. Canadian records of the Black Buffalo (see text for details). 



most commonly found in the deeper, swifter, 
cleaner waters of larger rivers and does not move 
into flooded areas, remaining in the permanent 
river channels. 

Although Trautman (1957) indicated that the 
species probably occupied a habitat intermediate 
to the Bigmouth and Smallmouth Buffalos, the 
habitat requirements seem to be closer to those of 
the latter. Clay (1962) stated that the species is 
intolerant of pollution; it is usually found in the 
deeper water of larger rivers, but may be found in 
marginal lakes (Smith 1979). 

General Biology 

There is little information on the biology of the 
species, but feeding and reproductive habits are 
said to be similar to those of the other buffalos 
(Smith 1979). Growth rates for the species in a 
Kansas reservoir were reported by Green and 
Cross ( 1 956), and Carlandcr ( 1 969) provides some 
additional age and growth data. Like most suckers, 
buffalos spawn in the spring after runoff raises 
stream levels. Thousands of eggs [up to 400 000 in 



larger fish, see Harlan and Speaker (1951)] are 
scattered over the bottom in shallow waters and 
abandoned. Hatching is assumed to occur in about 
10 days at 15°C [if similar to the Smallmouth 
Buffalo — see Smith (1979)] and young-of-the- 
year of all three species show up by mid-June {see 
Cross 1967; Smith 1979). Growth is rapid, young- 
of-the-year attaining 2 to 3 cm at the end of the first 
summer. They are sexually mature by age 3 at 22.5 
to 39 cm in length and 0.5 to 2 kg in weight (Harlan 
and Speaker 1951; Carlander 1969). By seven 
years, lengths of 100 cm or more may be achieved 
and fish of up to 24 years of age are known 
(Carlander 1969). One Black Buffalo taken in 
Kansas was 104.1 cm long and weighed 12.7 kg 
(Cross 1967). 

The species is benthic, feeding on plankton, 
insect larvae and vegetation; snails and small 
molluscs may also be important food items 
(Harlan and Speaker 1951). The species 
apparently hybridizes with Bigmouth and 
Smallmouth Buffalos where they are sympatric 
(Shute 1980). 



1990 



HOUSTON: Status of the Black Buffalo 



101 







Figure 3. Native distribution of the Black Buffalo, Ictiobus niger (adapted from Shute 1980). 



Limiting Factors 

No specific limiting factors have been indicated, 
except that the Black Buffalo has been reported as 
being intolerant of pollution (Clay 1962). The 
species is of Uttle or no commercial interest and is 
relatively rare compared to other buffalos (Smith 
1979). It is often taken by anglers using worms or 
doughballs for bait (Cross 1 967) but is not a sought 
after species (Harlan and Speaker 1951). 

Like the Smallmouth Buffalo, it seems less 
tolerant of turbidity (Trautman 1957) and may be 
limited by availability of suitable habitat and by 
damming of rivers for hydroelectric developments 
etc. It apparently does adapt to life in impound- 
ments (Cross 1967). They may also be hmited by 
competition with Carp (Cross 1967). The 
Canadian distribution may be limited by available 
habitat as the United States distribution has been 
shown to be sporadic, presence of the species being 
related to suitable habitat and water quality (Cross 
1967). 



Special Signiflcance of the Species 

Although of no commercial or sport interest, the 
species, like other buffalos, grows fast, does well in 
impoundments, and could be utilized as a source of 
protein. Its presence in the lower Great Lakes 
could be indicative of changing water quality. The 
extent of the Canadian distribution requires 
clarification. 

Evaluation 

The lower Great Lakes would appear to be the 
northern fringe of the native range of the species. 
Although locally abundant in some locations within 
the United States range, it is relatively rare as 
opposed to the other buffalos and has apparently 
declined or been extirpated in some parts of its 
range. United States and Canadian records for the 
species in Lake Erie are few and the species should 
be considered rare and vulnerable in Canada. 
Although only recently discovered in Canadian 
waters, there is no reason to suppose its presence in 



102 



The Canadian Field-Naturalist 



Vol. 104 



western Lake Erie is recent. Earlier United States 
records for the lake are known and the species is not 
easily identified and can be confused with the 
Smallmouth Buffalo, and younger fish of all these 
species are difficult to distinguish. 

Acknowledgments 

This report was made possible through the 
support of the Department of Fisheries and Oceans, 
World Wildlife Fund Canada and the Department 
of the Environment. 

Literature Cited 

Carlander, K. D. 1969. Handbook of freshwater fishery 

biology. Volume 1. Iowa State University Press, Ames, 

Iowa. 
Clay, W. M. 1962. A field manual of Kentucky fishes. 

Kentucky Department of Fish and Wildlife Resources, 

Frankfort, Kentucky. 
Cross, F. R. 1967. Handbook of fishes of Kansas. 

University of Kansas Museum of Natural History 

Miscellaneous Publication Number 45. 
Crossman, E. J., and S. J. Nepszy. 1979. First Canadian 

record of a Black Buffalo (Osteichthyes: Catostomidae). 

Canadian Field-Naturalist 93(3): 904-305. 



Green, J. K., and F. R. Cross. 1996. Fishes of El 

Dorado City Lake, Butler County, Kansas. Kansas 

Academy of Science Transactions 59: 958-963. 
Harlan, J. R., and E. B. Speaker. 1951. Iowa fish and 

fishing. Second edition. Iowa State Conservation 

Commission, Ames, Iowa. 
Hubbs, C. L., and K. F. Lagler. 1958. Fishes of the 

Great Lakes region. University of Michigan Press, 

Ann Arbour, Michigan. 
Johnson, J. E. 1978. Protected fishes of the United 

States and Canada. American Fisheries Society, 

Bethesda, Maryland. 
Moore, G. A. 1968. Fishes. Pages 21 -165 /« Vertebrates 

of the United States. Edited by 'W. F. Blair, A. P. Blair, 

P. Broadleob, F. R. Cagle, and F. A. Moore. McGraw 

Hill, New York, New York. 
Shute, J. R. 1980. Ictiobus niger. Black buffalo. Page 

406 in Atlas of freshwater fishes of North America. 

Edited by D. S. Lee, C. R. Gilbert, C. H. Hocutt, R. E. 

Jenkins, D. E. McAllister, and J. R. Stauffer Jr. North 

Carolina State Museum of Natural History, Biological 

Survey Publication Number 1980-12. 
Trautman, M. B. 1957. The fishes of Ohio. Ohio State 

University Press, Columbia, Ohio. 

Accepted 10 October 1989 



Status of the Golden Redhorse, Moxostoma erythrurum, 
in Canada* 



Cheryl D. Goodchild 

2168 Harcourt Crescent, Mississauga, Ontario L4Y IWl 

Goodchild, Cheryl D. 1990. Status of the Golden Redhorse, Moxostoma ervthrurum, in Canada. Canadian 
Field-Naturalist 1 04( 1 ): 1 03- 1 1 1 . 

The Golden Redhorse, Moxostoma erythrurum, is known from the Lake Huron, Lake St. Clair, and Lake Erie 
drainages in southwestern Ontario. Recent records from the Niagara River in Ontario and the Red River in Manitoba, 
indicate that its range is either expanding or more extensive than had previously been thought. This species is difficult 
to identify and is often overlooked or misidentified. Three other species of Moxostoma are either threatened or rare in 
Canada. Canadian populations are contiguous with those in the United States, but are at the northern limit of the 
range of the species. Little is knownof its biology in Canada, but it does not appear to beingreatjeopardy atthistime. 

Le suceur dore, Moxostoma erythrurum, frequente les bassins des lacs Huron, Sainte-Claire et Erie dans le sud-ouest 
de rOntario. Des donnees recentes, recueillies dans la riviere Niagara en Ontario ainsi que dans la riviere Rouge au 
Manitoba, indiquent que son aire de repartition est soit en expansion, soit plus etendue qu'on ne le croyait auparavant. 
L'espece est difficile a identifier et passe souvent inaper9ue ou est confondue avec une autre. Trois autres especes de 
Moxostoma sont soient menacees, soient rares au Canada. Les populations canadiennes, qui sent voisines des 
populations americaines, sont cependant a la Hmite septentrionale de I'aire de repartition du suceur dore. On connait 
peu de choses sur la biologic de l'espece au Canada, mais elle ne semble pas particulierement menacee a I'heure actuelle. 

Key Words: Golden Redhorse, Moxostoma erythrurum, suceur dore, Catostomidae, suckers, Moxostoma, redhorses, 
status, Canada. 



The Golden Redhorse, Moxostoma erythrurum 
(Rafinesque 1818), is generally gray to bronze or 
ohve coloured shading to white on the belly, with a 
golden cast in large specimens or a silvery cast in 
younger fish (Figure 1). Anal and paired fins are 
yellow to reddish orange. Trautman (1981) 
suggests that fishes taken from polluted waters 
may have all fins orange or reddish-orange. Adults 
are generally 280 to 450 mm (11 to 18 in) long, 
weighing up to 0.9 kg (2 lb). A 660 mm (26 in) long 
specimen was reported from Lake Erie (Trautman 
1981). 

This species is difficult to distinguish from other 
members of the genus, particularly the Black 
Redhorse, Moxostoma duquesnei, and the Silver 
Redhorse, Moxostoma anisurum. Kott (1979) 
found the most useful character to differentiate 
Golden Redhorse from Black Redhorse in 
southwestern Ontario, is the lateral line scale count 
(LLS), which ranges from 39 to 44 in Golden 
Redhorse and from 45 to 48 in Black Redhorse. 
Brown (1984) found the character (LLS) was 
highly variable and age dependent in Indiana and 
suggested that it was taxonomically limited. The 
obtuse angle formed by the lower lips is suggested 
as the best character for discriminating Golden 
Redhorse (Scott and Grossman 1973; Brown 



1984). Franzin et al. (1986) used body colouration, 
dorsal fin shape, and lower lip morphology in 
combination to separate three species of 
Moxostoma. 

Although formerly thought to be present only in 
southwestern Ontario, it has recently been 
reported from the Red River, Manitoba (Franzin 
et al. 1986) and the Lake of the Woods, in 
northwestern Ontario (V. ' Macins, Ontario 
Ministry of Natural Resources, Kenora, Ontario; 
personal communication). 

Distribution 

North America: The Golden Redhorse is 
restricted to the freshwaters of eastern North 
America (Figure 2). It occurs south from Lake 
Ontario tributaries of western New York State, 
west of the mountains, through western Pennsyl- 
vania and West Virginia to Tennessee and the 
north of Georgia, Alabama and Mississippi, west 
through much of northwestern Arkansas, eastern 
Oklahoma and extreme northern Texas, north 
through southeastern Kansas, through the eastern 
parts of the states from Nebraska to North 
Dakota, east across central Minnesota, Wisconsin, 
Michigan, and into Ontario (Scott and Grossman 
1973). 



*Reviewed and accepted by COSEWIC 1 1 April 1989 — no designation required. 

103 



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The Canadian Field-Naturalist 



Vol. 104 




Figure L Drawing of Golden Redhorse, Moxostoma erythrurum [drawing by Anker 
Odum, from Scott and Grossman (1973) by permission]. 



Lee et al. (1980) include the Upper Red River of 
the North, the Lake of the Woods drainage, and 
the Roanoke and James drainages on the Atlantic 
slope. Recently, it has also been found (perhaps 
introduced) in the Potomac drainage. The 
Potomac records apparently reflect a new range 
extension since it is absent from most Atlantic 
coast streams (Cooper 1983). The range of the 
Golden Redhorse may be expanding westward as 
well. In Kansas, Clarke and Clarke (1984) have 
reported it from the Arkansas River 160 km (100 
miles) west of previously known localities. 

Canada: Until recently, in Canada, the Golden 
Redhorse was believed to be restricted to the 
drainages of Lake Erie, Lake St. Clair and 
southern Lake Huron, in southwestern Ontario. 
Golden Redhorse have been collected from 125 
different locations in southwestern Ontario 
(Figure 3). The majority of the collections, 
however, have been made from a few major rivers 
and streams as follows: Catfish Creek, Grand 
River, Thames River, Nith River, Saugeen River, 
Sydenham River, Maitland River, and Nanticoke 
Creek. 

Scott and Grossman (1973) state that it was 
present in the Lake Erie system as far east as the 
Grand River, but not apparently in the Niagara 
River. More recent evidence suggests it may also be 
present in the Niagara River drainage in south- 
central Ontario. A single specimen was collected at 
the mouth of Miller Creek, near its confluence with 
the upper Niagara River in 1980 [Royal Ontario 
Museum, Toronto: ROM 37372]. This is perhaps 
not surprising as in New York State it is known 
from many streams draining into the Niagara 
River as well as in tributaries to Lake Ontario 
(Smith 1985). 

Dispersal into southwestern Ontario probably 
occurred before the arrival of man and was 
probably limited by temperature and suitable 
habitat (N. Mandrak, Department of Zoology, 
University of Toronto, Toronto, Ontario; personal 
communication). Postglacial dispersal into 



southwestern Ontario probably occurred 
through the Fort Wayne outlet of glacial Lake 
Maumee from a Mississippian refugium. Golden 
Redhorse occur in streams flowing into Lake 
Erie, Lake St. Clair and Lake Huron in Ontario. 
Niagara Falls, at Lewiston, was probably a 
barrier in both directions by approximately 
12 400 B.P. (Calkin and Feenstra 1985) and 
restricted it from moving eastward into the Lake 
Ontario drainage. The expansion of the 
populations in New York State, however, 
probably occurred due to migration through the 
Gennessee River on a fairly recent basis, since 
construction of the canals in the Lake Erie 
watershed provides reasonably free access (N. 
Mandrak, personal communication). 

Most recently, Moxostoma erythrurum has 
been reported from the Red River, Manitoba. 
Franzin et al. (1986) collected four adults about 1.5 
km downstream from St. Andrew's lock and dam 
near Lockport, Manitoba on 9 October 1981. An 
unconfirmed report of four individuals from the 
Lake of the Woods area of Ontario (Dechtiar 1 972) 
appears less doubtful with the recent collections in 
Manitoba and reports that Golden Redhorse occur 
in both the upper Red River (Lee et al. 1980) and 
Sheyenne River tributary in North Dakota 
(Peterka 1978). The Red River is part of the 
Hudson Bay Drainage (as is the Lake of the Woods 
area) and represents a significant extension of the 
known range of the Golden Redhorse (Figure 4). 

Moxostoma erythrurum may have dispersed 
into the Hudson Bay watershed later than glacial 
Lake Agassiz. It probably invaded from the 
Mississippi River into the Red River after the 
recession of ice floes due to limiting water 
temperature (N. Mandrak, personal communica- 
tion). Recent dispersal into the Red River may be 
restricted by a series of dams in North Dakota and 
Manitoba (Grossman and McAllister 1986) or it 
may be possible through the headwaters of the 
Minnesota and Red rivers during floods (Stewart 
and Lindsey 1983). 



1990 



GOODCHILD: STATUS OF THE GOLDEN REDHORSE 



105 




-30" 



100° 75"- 

Figure 2. North American distribution of the Golden Redhorse, Moxostoma erythrurum 
[adapted from Lee et al. (1980)]. 



Protection 

No specific protection other than that afforded 
by habitat sections of the Federal Fisheries Act 
exists in Canada. Since 1985, COSEWIC has 
suggested possibly assigning a rare status 
designation (Campbell 1985). It was listed as rare 
in Nebraska and South Dakota (Miller 1972). In 
Johnson's (1987) American Fisheries Society 
(AFS) list of protected fishes of the United States, 
and Canada it was designated as of special concern 
in both Manitoba and South Dakota. 

Population Sizes and Trends 

The actual population size of the Golden 
Redhorse has not been assessed in Canada. In 
southwestern Ontario, Golden Redhorse have 
been collected more or less continuously since the 
1920s and continue to be found in small numbers 
to the present. Difficulty in identifying the 
different species of redhorses hampers attempts to 
assess population size. In the 1970s, a large area of 
southwestern Ontario was surveyed by the Ontario 
Ministry of Natural Resources (OMNR) as part of 
the Stream Inventory Program. Approximately 
50% of all catalogued collections of Golden 
Redhorse were collected by OMNR during this 
period. Only small specimens were collected for 



positive identification, larger fish were returned to 
the water and clumped generally as redhorse 
suckers (G. Goodchild, Fisheries Branch, Ontario 
Ministry of Natural Resources, Toronto, Ontario; 
personal communication). 

In northwestern Ontario, Golden Redhorse 
have apparently been collected for years in the 
oligotrophic part of the Lake of the Woods but in 
low numbers (V. Macins, personal communica- 
tion), whereas Silver Redhorse are unknown from 
the Lake of the Woods although present 
throughout the surrounding area. 

In the Red River, Manitoba, W. G. Franzin, 
Department of Fisheries and Oceans, Winnipeg, 
Manitoba; personal communication) suspects that 
the species is rare probably because of restrictions 
in the amount of preferred habitat. 

It is an encouraging sign that the range of the 
Golden Redhorse in Canada appears to be 
expanding. Perhaps this is only indicative of 
increased survey efforts or better ability to 
distinguish Golden Redhorses from other redhorse 
species. Scott and Grossman (1973) felt that it was 
one of the few redhorses whose range did not seem 
to have been limited by habitat change. 

In New York State, Golden Redhorses are 
thought to be increasing in abundance as it has 



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Figure 3. Distribution of the Golden Redhorse, Moxostoma erythrurum, in southern Ontario. 



been collected more frequently in recent years 
(Smith 1985). Generally, in the United States, it 
appears to be faring well, often stated to be the 
most abundant redhorse in many states (Clay 1962; 
Cross 1967; Eddy and Underbill 1974; Pflieger 
1975). In Ohio, the Golden Redhorse has been the 
most widely distributed and most abundant species 
of Moxotoma since 1920 (Trautman 1981). It is, 
however, intolerant of pollutants, continuous 
turbidity and rapid siltation. 

Habitat 

The Golden Redhorse does not have particularly 
specialized habitat requirements and is, appar- 
ently, able to tolerate a fairly wide spectrum of 
habitat conditions. It is better adapted to river 
habitats than to lakes and is typically found in 
small to large streams and rivers with varied 
substrate (Lee et al. 1980). Avoidance of streams 
with high gradients is indicated by the paucity of 
records from such streams (Trautman 1981). It is 
associated with streams having large permanent 
pools. In Pennsylvania, Cooper ( 1983) found adult 
Golden Redhorse in slow deep runs of moderate- 
sized rivers. Kott et al. (1979) collected both 
Moxostoma duquesnei and Moxostoma erythru- 
rum in Ontario, al a site described as a stretch of 
swift-flowing shallow water over a gravel bottom 
ending in a one metre deep pool. 



Although frequently collected with the Black 
Redhorse, the Golden Redhorse prefers slightly 
warmer waters with less current and is more 
tolerant of turbidity and intermittent flow (Pflieger 
1975). In spite of its ability to withstand more 
siltation than most redhorse species, it has been 
eliminated from some streams by mine wastes 
(Clay 1962; Trautman 1981). According to 
Trautman (1981), it is seldom found in waters 
having an abundance of aquatic vegetation and 
only small numbers are present in larger lakes. On 
the other hand, Meyer (1962) remarks that the 
Shorthead Redhorse, Moxostoma macrolepido- 
tum, is commonly found in lakes near the northern 
margin of its range and this seems to be the case for 
the Golden Redhorse as well. It has been found in 
Lake Erie (Trautman 1981), Lake St. Clair [ROM 
1936, ROM 22892] and possibly the Lake of the 
Woods (V. Macins, personal communication) in 
the northern part of its range in Ontario. 

In southwestern Ontario where it is reasonably 
abundant, it is apparently able to withstand some 
variation in water quality. Fluctuating turbidity 
levels, oxygen levels and temperatures are reported 
for the Thames River system by McAllister ( 1 987), 
where the Golden Redhorse is commonly found. 
Rivers in southwestern Ontario are susceptible to 
these changes because it is a relatively flat, 
agricultural area. 



1990 



GOODCHILD: STATUS OF THE GOLDEN REDHORSE 



107 




Figure 4. Canadian distribution of the Golden Redhorse, Moxostoma erythrurum. 



Fluctuating water levels may have a detrimental 
effect on Golden Redhorse populations. Deacon 

(1961) found that in Arkansas, abundance of 
Golden Redhorse was directly related to stream 
conditions affected by drought, becoming scarce in 
the lower mainstream when flow increased. Meyer 

(1962) suggests that based on evidence presented 
by Hall and Jenkins (1953) and Finnell et al. 
(1956), the formation of reservoirs on rivers could 
have adverse effects on redhorse populations. 

One factor involved in maintenance of Golden 
Redhorse populations may be the accessibility of 
smaller streams for spawning and nursery habitat 
(Harlan and Speaker 1956). Trautman(1981) also 
reports that Golden Redhorse ascend the smaller 
streams in spring. Yet, in a study of the seasonal 
movement of Golden and Black redhorses in Ohio, 
no significant differences were found between 
average distances moved in the spring as compared 
to the fall, nor were any mass seasonal movement 
patterns exhibited (Smith 1978). 

Young Golden Redhorses have been described 
from varying habitats. Meyer (1962) and 
Larrimore et al. (1952) found young fish 
inhabitating slow-moving waters over soft- 
bottoms in areas near overhanging river banks. 



whereas Cooper (1983) found juveniles on riffle 
margins in moderate current. In contrast, Martin 
and Campbell (1953) found that, in Missouri, 
young Golden Redhorse inhabited the deeper, 
faster waters of streams near riffle areas. 

The complexities of proper identification of 
Moxostoma erythrurum, combined with difficul- 
ties in determining optimum habitat, precludes 
making definitive statements regarding protection 
of suitable habitat in Canada. 

General Biology 

Reproductive Capability: No studies on the 
biology of this species in Canada have been 
reported. Descriptions of the reproduction of the 
Golden Redhorse in the United States are available 
but often conflicting. 

Age and Growth: Maturity is reached between 
three and five, most commonly by age four (Meyer 
1962; Cross 1967; Lee et al. 1980; Smith 1985). 
Although the oldest age attained is generally 
reported to be seven years, Curry and Spacie 
(1984) suggest that recent work on annulus 
formation and age determination in White Suckers 
{Catostomus catostomus) by Quinn and Ross 
(1982) may cast doubt on reUability of aging 



108 



The Canadian Field-Naturalist 



Vol. 104 



catostomids older than age five, and perhaps they 
cease to form annuli after age seven. For age seven 
and older Black Redhorse, Bowman (1970) found 
that the percentage of scales with annuli completed 
by May decreased with increasing age, but he was 
able to age fish up to nine years. 

Although Trautman (1979) gave a maximum 
size from Lake Erie as 660 mm (26 in) in length, 
and 2 kg (4.5 lb) in weight, he stated that the usual 
length was 279 to 457 mm (11 to 18 in). Meyer 
(1962) found they reached an average length of 84 
mm (3.3 in) by age one and 488 mm (19.2 in) by age 
seven, whereas in Oklahoma they reached a 
maximum size of 625 mm (24.6 in) [Scott and 
Grossman 1973]. In Missouri, they averaged 79 
mm (9. 1 in) at the end of their first year, and about 
150 mm (9.9 in), 218 mm, 272 mm, 310 mm at the 
end of each succeeding year, with an average life 
span of six to seven years. Few fish over 1 1 years of 
age were reported. Meyer (1962) found no 
difference in growth rates between sexes and found 
that most of the growth occurred in late July, 
August and early September. 

Spawning: Golden Redhorses spawn in spring, 
usually in May or early June and later than other 
redhorses in the same area, (Scott and Grossman 
1973). In Iowa, spawning occurs after water 
temperatures rise to greater than 16°G (60° F) 
[Meyer 1962], but in Kansas, temperatures of 
greater than 22° G (70° F) have been noted (Gross 
1967). Both fall within the range of 17° to 22° G 
given by Lee et al. (1980). 

Spawning lasts for only a short period. Males 
are generally observed at the spawning sites prior 
to the arrival of the females, apparently defending 
territories. Sex ratios were found to be 75 males to 
37 females indicating the brevity of the females stay 
(Mayer 1962). Sex ratios at other times of the year 
were 1:1. Non-adhesive eggs are broadcast over 
gravel riffles and abandoned (Gooper 1983). 
Meyer ( 1 962) estimated that the average number of 
eggs per female is 6100 to 25 390, but estimates ran 
as high as 35 000 in Ohio (Garlander 1950). 

Depth or flow may be a critical factor in the 
selection of spawning sites as suggested by Gurry 
and Spacie (1984). They observed that spawning 
did not take place until stream levels had dropped 
and velocity decreased. Smith (1977) reported 
stream velocities of 0.9 to 1.2 m.s' during 
spawning of Golden Redhorse in Glear Greek, 
Iowa. Reproductively active males usually occupy 
shallow water with moderate current (Gross 1967). 
Spawning in shallow water up to 0.6 m deep is 
reported by various authors (Reighard 1920; 
Meyer 1962; Eddy and Underbill 1974). 

Species Movement: One of the most controver- 
sial aspects of the biology of the Golden Redhorse 



is the disagreement over whether or not the species 
migrates to any appreciable degree. Scott and 
Grossman (1973) describe it as being sedentary, 
remaining in or near the same pool with occasional 
marked downstream movements. Short migra- 
tions upstream have also been frequently observed 
(Gross 1967; Bowman 1970; Smith 1977; Gooper 
1983). Trautman (1981) states that the Golden 
Redhorse has a highly migratory nature. Yet, 
Meyer (1962) and Smith (1978) found no evidence 
for movement. Individuals marked by Deacon 
(1961) in Kansas during stable water levels were 
recaptured in areas of original capture and release. 
Gross ( 1 967) also found the Golden Redhorse to be 
sedentary except during periods of fluctuating 
water levels. Gerking (1953) states that the Golden 
Redhorse does not ascend small streams in Indiana 
unless they are near their home territory. 

Studies undertaken by Gurry and Spacie (1984) 
may explain conflicting observations on spawning 
movements that have been reported. They found 
that there was an age/ size differential in the use of 
tributaries. Larger adults do not apparently 
expend the energy to move upstream where 
smaller/ younger adults are more common. 
Segregation of spawning groups by size acts to 
distribute the competitive advantage for the 
Golden Redhorse. 

Further studies might be undertaken to 
determine if the degree of migration is related to 
preferred habitat regarding depth and rate of flow 
for various age/ size groups. Stream alterations 
that affect temperature, depth, flow regimes, and 
fish movement would tend to have an impact on 
populations. For this reason, the construction of 
dams or reservoirs could have debilitating effects 
on populations. 

Adaptability/ Behaviour: The Golden Redhorse 
feeds primarily on small molluscs, crustaceans, 
insects, detritus and algae (Lee et al. 1980). Gooper 
( 1 983) describes it as a benthic feeder. Results of a 
Iowa study to determine principal foods for 
redhorse above four inches long are listed: 
immature chironomids (91%), immature Ephe- 
meroptera (62%), and immature Trichoptera 
(18%) [Meyer 1962]. No differences in food habits 
were found between Golden Redhorse, Silver 
Redhorse and Shorthead Redhorse. Significant 
differences in the major groups of food consumed 
by Golden Redhorse were found between age 
groups and different populations (Brown 1984). 

The following group of parasites were Hsted by 
Hoffman (1967): Protozoa, Trematoda, Nemat- 
oda, Acanthocephala, leeches. 

Since the Golden Redhorse is still considered 
common in most areas of its range in the United 
States, it is probably reasonably tolerant of human 
disturbance, its continued existence in Ganada 



1990 



GOODCHILD: STATUS OF THE GOLDEN REDHORSE 



109 



may be more tenuous due to being at the northern 
extent of its range. 

Limiting Factors 

There is no evidence to suggest that the Golden 
Redhorse populations are declining in Canada. It 
is difficult to determine whether the broader 
distribution of the Golden Redhorse has resulted 
from true range extensions or because of 
asystematic sampling and difficulties in identifying 
species of redhorse suckers. 

Although apparently more tolerant to habitat 
alterations than the Black Redhorse, it does seem 
to be susceptible to changes that affect stream 
depth or velocity of flow. Any damming or 
reservoir development on Golden Redhorse 
streams would have potentially adverse effects. In 
Ontario, several dams on the Grand and Thames 
rivers have reduced the amount of suitable habitat 
for the Black Redhorse (Parker and Kott 1987). 
Doubtless, these dams have had detrimental effects 
on Golden Redhorse populations as well. 

The area of southwestern Ontario occupied by 
the Golden Redhorse is rural, characterized by 
large areas cleared for agricultural use. The degree 
of impact from agricultural practices — pesticides, 
siltation, stream channelizations, irrigation, and 
reservoirs is uncertain, but bound to have a 
negative effect on Golden Redhorse populations. 

Angling probably has a negligible effect on 
Golden Redhorse populations. The Golden 
Redhorse is not sought after in Canada, but it is 
taken incidentally on hook and line with natural 
bait. 

There is some recreational fishing for the Golden 
Redhorse in the United States. In Kansas, the 
Golden Redhorse had long supported a sport 
fishery (Cross 1967). Due to local abundance in 
Pennsylvania, it is often taken by anglers and often 
preferred over other suckers for food (Cooper 
1983). In other states such as Iowa, it is not a 
favoured species but they are taken occasionally 
from streams (Harlan and Speaker 1956). 

Scott and Grossman (1973) considered redhorse 
suckers to be commercial species of some 
importance in some areas. Since all species of 
redhorses are generally marketed as suckers or 
mullet, it is difficult to appraise the contribution to 
the catch of one species. However, they felt that 
both size and stream habitat precluded any 
significant entry into the commercial catch by the 
Golden Redhorse. 

Both young and adults of this species probably 
do not compete for food or space with more valued 
species (Scott and Grossman 1973). Meyer (1962) 
found no difference in food habits between the 
Golden, Silver, and Shorthead redhorse. Although 
these species often co-exist in the same streams, 



slight variations in the preferred areas occupied 
may diminish the amount of competition for food. 
Except for young fish. Golden Redhorses are 
not likely subject to significant predation. 
Predation on adults is probably limited by their 
size and the faunal composition of their habitat 
(Scott and Grossman 1973). 

Special Significance of the Species 

The Golden Redhorse, belongs to the genus 
Moxostoma, one of the most perplexing groups of 
North American fishes. Despite numerous species 
and large size, very little information is available 
on life history. The uncertain systematic position 
of the species is due to the few interspecific 
differences exhibited in meristic features (Scott 
and Grossman 1973). 

In Canada alone, there are seven species of 
Moxostoma. Because of problems in distinguishing 
between these species, capture records are 
unreliable. For this reason, and due to sporadic 
sampling, it is difficult to determine whether 
populations are stable or declining. Three species of 
redhorse are considered threatened or rare in 
Canada. Moxostoma carinatum, the River 
Redhorse, is considered rare; the Black Redhorse 
and Moxostoma hubbsi, the Copper Redhorse, are 
considered threatened (Parker and McKee 1984; 
McAlhster et al. 1985; Parker and Kott 1987). 
Canadian populations are apparently contiguous 
with those in the United States, but are at the 
northern limit of the range of the Golden Redhorse. 
The species is of some minor importance as a bait 
fish and to sport and commercial fishermen. 

Evaluation 

Whether recent records updating the distribu- 
tion of the Golden Redhorse are a result of recent 
range extensions or of more intensive samphng is 
difficult to judge. No population studies have been 
done in Canada but collection records indicate it is 
still present throughout most of its range in 
southwestern Ontario. Status of populations in 
Manitoba have not been ascertained. While 
populations of the Golden Redhorse do not appear 
to be under immediate threat, the species is 
vulnerable to changes in depth, rate of flow, 
excessive turbidity and pollution. 

Until more information is available on popula- 
tion trends, the status of the species in Canada is 
difficult to ascertain. However, existing populations 
appear to be stable and there is no requirement for 
COSEWIC status at this time. 

Acknowledgments 

This report has been funded by the World 
Wildlife Fund, Canada. Sincere thanks to R. R. 
Campbell, Fisheries and Oceans Canada, for his 



110 



The Canadian Field-Naturalist 



Vol. 104 



support in the preparation of this report. The assist- 
ance of D. E. McAlUster of the National Museum of 
Natural Sciences (now Canadian Museum of 
Nature), E. Holm of the Royal Ontario Museum, 
G. E. Gale, and G. A. Goodchild, of the Ontario 
Ministry of Natural Resources, in providing access 
to records and reports was, invaluable. 

W. R. Scott, Huntsman Marine Laboratory and 
E. J. Grossman of the Royal Ontario Museum 
gave permission to use the drawing of Moxostoma 
erythrurum from the Freshwater Fishes of 
Canada, Bulletin 184, Fisheries Research Board of 
Canada (1973). 

The author is grateful to P. McKee, Beak 
Consultants, for providing the southwestern 
Ontario "base" map, and to N. E. Mandrak, 
University of Toronto, for his insight into the 
distribution of this species. The assistance of E. J. 
Grossman, of the Royal Ontario Museum, V. 
Macins, of the Ontario Ministry of Natural 
Resources, and W. G. Franzin, Fisheries and 
Oceans Canada, is gratefully acknowledged. 

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Redhorse, Moxostoma duquesnei (LeSueur), in 
Missouri. Transactions of the American Fisheries 
Society 99(3): 546-559. 

Brown, R. A. 1984. Comparative life histories of some 
species of redhorse, Subgenus Moxostoma, genus 
Moxostoma. Ph.D. dissertation, Indiana State 
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Calkin, P. E., and B. H. Feenstra. 1985. Evolution of the 
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Campbell, R. R. Editor. 1987. Rare and endangered 
fishes and marine mammals of Canada. COSEWIC 
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Carlander, K. D. 1950. Handbook of freshwater fishery 
biology. W. C. Brown, Dubuque, Iowa. 281 pages. 

Clarke, R. F., and J. W. Clarke. 1984. New county 
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Clay, W. M. 1962. A field manual of Kentucky fishes. 
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Frankfort, Kentucky. 147 pages. 

Cooper, E. L. 1983. Fishes of Pennsylvania and the 
northeastern United States. Pennsylvania State 
University Press, University Park, Pennsylvania. 243 
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Cross, F. B. 1967. Handbook of fishes of Kansas. 
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( rossman, E. J., and O. E. McAllister. 1986. Zoogeo- 
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Wiley. J. Wiley and Sons, New York, New York. 



Curry, K. D., and A. Spacie. 1984. Differential use of 
stream habitat by spawning catostomids. American 
Midland Naturalist 1 1 1 (2): 267-279. 

Deacon, J. E. 1961. Fish populations, following a 
drought, in the Neosho and Marias des Cygnes Rivers 
in Kansas. University of Kansas, Museum of Natural 
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Dechtiar, A. O. 1972. Parasites of fish from Lake of 
the Woods, Ontario. Journal of the Fisheries 
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Eddy, S., and S. C. Underhill. 1974. Northern fishes 
with special reference to the upper Mississippi Valley. 
Third edition. University of Minnesota Press. 
Minneapolis, Minnesota. 414 pages. 

Finnell, J. C, R. M. Jenkins, and G.E. 
Hall. 1956. The fishery resources of the Little River 
system, McCurtain County, Oklahoma. Oklahoma 
Fisheries Research Laboratory Report 55. 82 pages. 

Franzin, W. G., B. R. Parker, and S. M. Harbicht. 
1986. First record of the Golden Redhorse, 
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Manitoba. Canadian-Field Naturalist 100(2): 
270-271. 

Gerking, S. D. 1953. Evidence for the concepts of 
home range and territory in stream fishes. Ecology 
34(2): 347-965. 

Hall, G. E., and R. M. Jenkins. 1953. Continued 
fisheries investigations of Tenkiller Reservoir, 
Oklahoma, during its first year of impoundment, 
1953. Oklahoma Fisheries Research Laboratory 
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Harlan, J. R., and E. B. Speaker. 1956. Iowa fish and 
fishing. Third edition. Iowa State Conservation 
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Hoffman, G. L. 1967. Parasites of North American 
freshwater fishes. University of California Press, Los 
Angeles, California. 486 pages. 

Hubbs, C. L., and K. F. Lagler. 1967. Fishes of the 
Great Lakes Region. University of Michigan Press, 
Ann Arbor, Michigan. 213 pages. 

Jenkins, R. E. 1970. Systematic studies of the 
catostomid fish tribe Moxostomatini. Ph.D thesis, 
Cornell University, Ithaca, New York. 818 pages. 

Johnson, J. E. 1987. Protected fishes of the United 
States and Canada. American Fisheries Society, 
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Kott, E., R. E. Jenkins, and G. Humphreys. 1979. 
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toma duquesnei from Ontario. Canadian 
Field-Naturalist 93(1); 63-66. 

Larimore, R. W., Q. H. Pickering, and L. Durham. 
1952. An inventory of the fishes of Jordan Creek, 
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Lee, D. S., C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, 
D.E. McAllister, and J. R.Stauffer, Jr. 1980. Atlas 
of North American freshwater fishes. North Carolina 
State Museum of Natural History, Biological Surveys 
Publication 1980-12. 867 pages. 

Martin, R. O., and R. S. Campbell. 1953. The small 
fishes of Black River and Clearwater Lake, Missouri. 
University of Missouri Studies 26: 49 66. 

McAllister, D. E. 1987. Status of the Central Stone- 
roller, Campostoma anomalum, in Canada. Cana- 
dian licld Naturalist 101(2): 213 218. 



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McAllister, D. E., R. J. Parker, and P. M. McKee. 

1985. Rare, endangered and extinct fishes in Canada. 

National Museum of Natural Sciences Syllogeus 

Number 54: 1-192. 
Meyer, W. H. 1962. Life history of three species of 

redhorse (Moxostoma) in the Des Moines River, Iowa. 

Transactions of the American Fisheries Society 91(4): 

412-419. 
Miller, R. R. 1972. Threatened freshwater fishes of the 

United States. Transactions of the American Fisheries 

Society 101(2): 239-252. 
Parker, B., and E. Kott. 1987. Status report on the Black 

Redhorse (Moxostoma duquesnei) in Canada. Report 

to the Committee on the Status of Endangered Wildlife 

in Canada. Canadian Wildlife Service, Ottawa, Ontario. 

19 pages. 
Parker, B., and P. McKee. 1984. Status of the River 

Redhorse, Moxostoma carinatum, in Canada. 

Canadian Field-Naturalist 98(1): 110-114. 
Pflieger, W. L. 1975. The fishes of Missouri. Missouri 

Department of Conservation, Columbia, Missouri. 343 

pages. 
Purkett, C. A., Jr. 1958. Growth rates of Missouri stream 

fishes. Missouri Conservation Commission, D.-J. 

Report Number 1. 46 pages. 
Quinn, S. P., and M. R. Ross. 1982. Annulus formation 

by white suckers and the reliability of pectoral fin rays 

for aging them. North American Journal of Fish 

Management 2: 204-208. 
Reighard, J. 1920. The breeding behaviour of the suckers 

and minnows. Biological Bulletin 38(1): 1-32. 



Scott, W.B. 1967. Freshwaterfishes of eastern Canada. 
Second edition. University of Toronto Press, Toronto, 
Ontario. 137 pages. 

Scott, W. B., and E.J. Grossman. 1973. Freshwater 
fishes of Canada. Fisheries Research Board of Canada 
Bulletin 184: 1-966. 

Smith, C. A. 1977. The biology of three species of 
Moxostoma in Clear Creek, Hocking and Fairfield 
counties, Ohio, with emphasis on the Golden 
Redhorse, M. erythrurum (Rafinesque). Ph.D. thesis, 
Ohio State University, Columbus, Ohio. 158 pages. 

Smith, C. G. 1978. The seasonal movement of Golden 
and Black Redhorses {Moxostoma erythrurum and M. 
duquesnei) in Clear Creek, Fairfield and Hocking 
Counties, Ohio. Ohio Journal of Science 78 
(supplement): 13. 

Smith, C. L. 1985. The inland fishes of New York State. 
New York State Department of Environmental 
Conservation, Albany, New York. 522 pages. 

Stewart, K. W., and C. C. Lindsey. 1983. Postglacial 
dispersal of lower vertebrates in the Lake Agassiz 
region. Pages 391-419 in Glacial Lake Agassiz. Edited 
by J. Teller and Lee Clayton. Geographic Association 
of Canada Special Paper 26. 

Trautman, M. B. 1981. The fishes of Ohio with 
illustrated keys. Revised edition. Ohio State University 
Press, Columbus, Ohio. 782 pages. 

Accepted 10 October 1989 



Status of Ball's Porpoise, Phocoenoides dalli, in Canada* 

Thomas A. Jefferson 

Marine Mammal Research Program, Texas A & M University, P.O. Box 1675, Galveston, Texas 77553-1675 

Jefferson, Thomas A. 1990. Status of Call's Porpoise, Phocoenoides dalli, in Canada. Canadian Field-Naturalist 
104(1): 112-116. 

Dall's Porpoise, Phocoenoides dalli, is one of the most commonly sighted cetaceans throughout its range in temperate 
waters of the North Pacific Ocean and surrounding seas. Dall's Porpoises are common both offshore and in deep 
inshore waters of British Columbia. There appear to be few serious conservation problems in the eastern Pacific at 
present, although little is known of the behaviour and ecology of this species. Probably the major threat facing this 
species in Canada is environmental contamination by such substances as organochlorines and heavy metals. Much 
more research is needed before these threats can be properly assessed but, in the meantime, a conservative approach to 
porpoise management is suggested. 

Le marsouin de Dall, Phocoenoides dalli, est I'un des cetaces les plus couramment apergus dans toute son aire de 
distribution en eaux temperees de I'ocean Pacifique nord et des mers viosines. Le marsouin de Dall est commun tant 
dans les eaux du large que dans les eaux cotieres profondes de la Colombie-Britannique. Actuellement, la conservation 
de cette espece presente peu de problemes importants dans Test du Pacifique, bien qu'on connaisse peu de chose de son 
comportement et de son ecologie. La contamination de I'environnement par, notamment, les organochlores et les 
metaux lourds constitue probablement le principal danger pouvant menacer I'espece au Canada. Une evaluation 
adequate de ce genre de dangers necessiterait des recherches beaucoup plus importantes et Ton propose, entre temps, 
de proceder avec prudence a la gestion de ce marsouin. 

Key Words: Dall's Porpoise, Phocoenoides dalli, cetaceans, British Columbia, status. 



Dall's Porpoise, Phocoenoides dalli {Trwt 1885), 
despite being a commonly-sighted species in the 
North Pacific, is rather poorly known in most 
aspects of its ecology and population biology. The 
published literature on this species has recently 
been reviewed (Jefferson 1988), and therefore this 
report focuses only on information relevant to its 
status in British Columbia. 

Dall's Porpoise is the largest member of the 
porpoise family (Phocoenidae), growing to lengths 
of about 220 cm and weights of 200 kg (Leather- 
wood et al. 1982). These animals are extremely 
robust, with small heads and small appendages 
(Figure 1). The wide-based triangular dorsal fin is 
slightly recurved at the tip, and the caudal 
peduncle is strongly keeled, especially in adult 
males (Jefferson 1990). 

The colour pattern is diagnostic. A ventrally- 
continuous white patch extends up on both flanks 
and forward to the level of the dorsal fin on the 
predominantly black body. White to light gray 
areas occur on the upper half to two-thirds of the 
dorsal fin, and the rear borders of the flukes. This 
"frosting" is not present on newborns. There 
appear to be several colour morphs, including 
gray, all-black, and all-white forms. A discrete 
population off the Pacific coast of Japan has a 
larger flank patch (this colour morph is called 
/rue/-typc, as opposed to dalli-typc; Kasuya 1982). 



Distribution 

Phocoenoides dalli is a North Pacific endemic, 
being found from northern Baja California, 
Mexico, north to the southern Chukchi Sea, and 
south to southern Japan (Figure 2). The species is 
only common between 32° N and 62° N in the 
eastern North Pacific (Nishiwaki 1967; Morejohn 
1979). 

Off the west coast of Canada, Dall's Porpoises 
are found mostly over the Continental Shelf and 
slope, but also more than 2400 km from shore 
(Pike and MacAskie 1969). They are seen year- 
round in the deeper inshore waters of British 
Columbia (Leatherwood et al. 1982). 

There are 15 published specimen records and 
over 300 sighting records from British Columbia 
(Cowan 1944; Scheffer 1949; Pike and MacAskie 
1969; Jefferson 1987; Baird et al. 1988) Off the 
coast of Canada, Dall's Porpoises are found 
mostly over the continental shelf and slope, but 
occasionally more than 2400 km from shore 
(Pike and MacAskie 1969). They are seen year- 
round in the deeper inshore waters off British 
Columbia, such as Hecate Strait, Laredo 
Channel, Queen Charlotte Sound, Goletas 
Channel, Queen Charlotte Strait, Johnstone 
Strait, and Strait of Juan de Fuca (Cowan 1944; 
Pike and MacAskie 1969; Leatherwood et al. 
1982; Jefferson 1987). 



■Reviewed and accepted by COSEWIC I I April 1989 no designation required. 

112 



1990 



JEFFERSON: STATUS OF DaLL'S FURi'UiSh 



113 




Figure 1. Adult Female Dall's Porpoise, Phocoenoides dalli (photograph by the 
author). 



Protection 

Dall's Porpoise is protected in Canadian 
waters under the 1982 Cetacean Protection 
Regulations of the Fisheries Act of Canada of 
1970. In United States waters, primary protection 
is provided by the Marine Mammal Protection 
Act of 1972. International protection measures 
include listing in Appendix II of the Convention 
on International Trade in Endangered Species of 
Wild Fauna and Flora (CITES), which regulates 
international trade. 

Population Size and Trends 

The population status of animals in the eastern 
North Pacific is not known, and there are no 
abundance estimates for British Columbia. 
Leatherwood et al. (1982) called Phocoenoides 
dalli the most abundant porpoise north of 
Vancouver Island. The entire North Pacific 
Ocean and Bering Sea population is estimated to 
be 1.4 to 2.8 million animals (Jones et al. 1987). 
The population trend is unknown, but because 
there is no evidence of major mortality, it is 
assumed to be relatively stable in the eastern 
Pacific. The western Pacific and Bering Sea 
stocks are heavily exploited and some may be at 
risk (Jones et al. 1987; Perrin 1* 



Habitat 

The species is widespread along the British 
Columbia coast, and is quite common both 
inshore and offshore. Cowan (1944) suggested 
that in inshore areas, Dall's Porpoises select 
open-ended channels with strong currents. The 
primary habitat is cool (<17°C), deep 
(> 180 m), continental shelf and slope waters 
(Jefferson 1988). Morejohn (1979) suggested that 
movements are mainly due to availability of prey, 
which consists of various species of squid and 
small schooling fishes. Many different prey 
species are known from throughout the range, 
and the Dall's Porpoise is thought to be an 
opportunistic feeder (Stroud et al. 1981), 



although the stomachs of four specimens from 
British Columbia contained only Pacific Herring, 
Clupea harengus (Cowan (1944). There is no 
evidence of significant habitat change in British 
Columbia. 

General Biology 

Reproductive Capability: Very little informa- 
tion is available on Dall's Porpoise reproduction 
in the eastern North Pacific. There is apparently a 
very strong summer calving peak (as in the rest of 
the range) in the months of June through August, 
and a smaller peak in March (Jefferson 1989). 

Most information comes from the western 
Pacific, where large-scale fishery interactions 
provide large samples for analysis. Dalli-type 
males become sexually mature at an age of 4 to 6 
years and a length of 180 to 186 cm, and females 



5«. 


-J 

Alaska 


P 




A 


Z^-^'--, 


/ 




yg 


^ 




anada ^o"- 
i USA 




140° 




120° V 



Figure 2. General distribution of the Dall's Porpoise 
in the eastern North Pacific. 



114 



The Canadian Field-Naturalist 



Vol. 104 



at 3.5 to 4.5 years and 174 to 177 cm (Kasuya and 
Shiraga 1985; Jones et al. 1987; Miyazaki 1987). 
Truei-iype animals become mature at body 
lengths about 12 to 17 cm greater than these 
(Kasuya 1978; Kasuya and Shiraga 1985). Most 
females appear to have an annual reproductive 
cycle (Kasuya and Jones 1984; Jones et al. 1987). 
Gestation lasts about 10 to 11 months (Jones et 
al. 1983); and the lactation period is unknown, 
but it is thought to be very short, perhaps 2 to 4 
months (Newby 1982). 

Species Movement: Dall's Porpoises in the 
eastern North Pacific, although present through- 
out their range year-round, tend to have inshore 
and southern shifts in abundance for the winter, 
and offshore and northern shifts for the summer 
(Leatherwood et al. 1982). They are present both 
inshore and offshore in British Columbia all year, 
although there seems to be an offshore shift in 
abundance for the summer (Pike and MacAskie 
1969; Leatherwood et al. 1982). 

Behaviour/ Adaptability: These animals are 
avid bow-riders, and are thought to be among the 
fastest swimming small cetaceans. When bow- 
riding or moving quickly they produce a 
distinctive rooster-tail splash. Porpoises not 
reacting to vessels often surface in a slow roll, 
with no splash. At these times, the deepened 
peduncle is visible above the surface. This type of 
surfacing seems to be predominant in inshore 
waters (Jefferson 1987; Miller 1988a). Dall's 
Porpoises rarely breach or indulge in other aerial 
behaviour. 

Porpoises are attracted to vessels in both 
inshore and offshore waters, especially to bow- 
ride, but they sometimes avoid vessels as well 
(Withrow et al. 1985) In particular, cow/calf 
pairs often avoid ships (Kasuya and Jones 1984). 

Groups are generally small (less than 10 
animals) and fluid, and are composed of very 
small subgroups which may aggregate at times, 
especially for feeding (Miller 1988b). Large 
aggregations of up to several thousand are 
occasionally seen (Scheffer 1950). 

The degree of habitat specialization has not 
been well studied in this species. Inshore calving 
areas have been proposed for both major study 
areas in Puget Sound, Washington and John- 
stone Strait, British Columbia (Jefferson 1987; 
Miller 1988a), but young calves are seen offshore 
as well, especially in the western Pacific (Kasuya 
and Jones 1984; Kasuya and Ogi 1987). Feeding 
presumably occurs throughout the range. 

Limiting Factors 

Killer Whales, Orcinus orca, and to a lesser 
extent, large sharks, are potential predators but 



the degree of predation is unknown (Pike and 
MacAskie 1969; Morejohn 1979; Newby 1982). 
Dall's Porpoises are rarely preyed upon by 
resident whales in British Columbia, but 
transient Killer Whales attack marine mammals 
(Bigg et al. 1987). There is little evidence that 
sharks regularly take healthy animals (Dall's 
Porpoises may swim too fast for most sharks), 
although one may speculate that they may weed- 
out sick or injured porpoises that are less able to 
avoid them. 

Parasite infestations are common and often 
extensive in this species (Walker 1975), but the 
role they play in natural mortality is unknown. 
Disease factors have not been adequately studied 
in Phocoenoides dalli. 

Human disturbance or disruption of activities 
is not thought to be a serious problem at present. 
As noted by Miller (1988a), these animals 
generally have complete control over encounters 
with boats, and it is indeed nearly impossible to 
follow them if they do not want to be followed. 
Heavy sustained vessel traffic in major feeding or 
breeding areas, however, could potentially 
disrupt activities or cause abandonment of the 
area. Future oil and gas exploration activities 
could pose a threat to this species, although the 
risk is still largely unknown (Geraci and St. 
Aubin 1980; W'ursig 1990). One can speculate 
that a major oil spill in a critical feeding area 
could be harmful. 

Although some incidental morality occurs due 
to fishing operations, no major fishery conflicts 
are known in Canada. Small numbers have been 
reported to be captured in gillnets and trawl nets 
(Everitt et al. 1979; Jefferson 1987; R. W. Baird, 
Victoria, British Columbia; personal communi- 
cation), and 58 were taken in 1987 in the 
experimental squid driftnet fishery, but this 
fishery has since been discontinued (Baird et al. 
1988). The species has a tendency to entangle in 
gillnets, however, and the possibility of 
undocumented problems exists. 

The largest potential threat to this species in 
Canadian waters may be environmental pollution 
by human activities. Little work has been done in 
the eastern Pacific, but Japanese scientists have 
recently explored levels of organochlorines and 
heavy metals in porpoises from the western 
Pacific. Small cetaceans were found to be poorly 
equipped to metabolize PCBs (Tanabe et al. 
1988). High PCB and DDE levels can result in 
decreased testosterone levels, possibly impairing 
reproduction (Subramanian et al. 1987b). 
Perhaps most disturbing was the discovery that 
excretion of pollutants by females occurs mainly 
through parturition and lactation (Subramanian 
et al. i987a). Thus, it is possible that young Dall's 



1990 



JEFFERSON: STATUS OF DALL'S PORPOISE 



115 



Porpoises in some areas are starting their lives 
with already high levels of environmental 
contaminants. 

Special Significance of the Species 

Dall's Porpoises are known to most people 
who frequent the deeper coastal waters of the 
western United States and British Columbia. 
Although they support no industry of their own, 
Dall's Porpoises often delight passengers on 
whale-watching and nature cruises along the west 
coast. Their bow-riding antics never cease to 
thrill those lucky enough to see them, and these 
cruises would certainly lose some of their appeal 
without them. 

The species normaly feeds on small schooling 
fishes and squids that are not highly prized by 
sport and commercial fishermen. Because it is an 
upper level carnivore that accumulates pollu- 
tants, it might be used as a biological indicator of 
the health of local ecosystems (Tanabe et al. 
1983). 

Evaluation 

There is no evidence that Dall's Porpoise 
populations in British Columbia are being 
depleted at present. Although several western 
Pacific populations may be depleted, Phoco- 
enoides dalli is considered to be common in 
Canada. However, there is a lack of information 
on potential threats, and until more is known, a 
conservative management approach should be 
taken. 

Acknowledgments 

Thanks to Robert Campbell, Chairman of the 
COSEWIC Fish and Marine Mammal Subcom- 
mittee, for advice and encouragement; and to 
Bernd Wursig, Pam Stacey, and Beth Miller for 
useful reviews of the manuscript. Robin Baird 
also reviewed the paper and provided a great deal 
of helpful information. This represents Contribu- 
tion No. 8 of the Marine Mammal Research 
Program, Texas A&M University at Galveston. 

Literature Cited 

Baird, R. W., K. M. Langelier, and P. J. Stacey. 198- 
8. Stranded whale and dolphin program of B.C. — 
1987 Report. British Columbia Veterinary Medical 
Association Wildlife Veterinary Report 1: 9-12. 

Bigg, M. A., G. M. Ellis, J. K. B. Ford, and K. C. 
Balcomb. 1987. Killer whales: A study of their 
identification, genealogy and natural history in 
British Columbia and Washington State. Phantom 
press and Publishers Inc., Nanaimo, British 
Columbia. 

Cowan, I. M. 1944. The Dall Porpoise, Phocoenoides 
dalli (True), of the northern North Pacific. Journal of 
Mammalogy 25: 295-306. 



Everitt, R. D., C. H. Fiscus, and R. L. Delong. 1979. 
Marine mammals of northern Puget Sound and the 
Strait of Juan de Fuca: A report on investigations 
November 1, 1977 — October 31, 1978. NOAA 
Technical Memorandum ERL MESA-41: 1-191. 

Geraci, J. R., and D. J. St. Aubin. 1980. Offshore 
petroleum resource development and marine 
mammals: A review and research recommendations. 
Marine Fisheries Review 42: 1-12. 

Jefferson, T. A. 1987. A study of the behavior of 
Dall's Porpoise, Phocoenoides dalli in the Johnstone 
Strait, British Columbia. Canadian Journal of 
Zoology 65: 736-744. 

Jefferson, T. A. 1988. Phocoenoides dalli. Mammal- 
ian Species 319: 1-7. 

Jefferson, T. A. 1989. Calving seasonality of Dall's 
Porpoise in the eastern North Pacific. Marine 
Mammal Science 5: 196-200. 

Jefferson, T. A. 1990. Sexual dimorphism and 
development of external features in Dall's porpoise 
Phocoenoides dalli. Fishery Bulletin (U.S.) 88: 
119-132. 

Jones, L. L., D. W. Rice, and A. A. Wolman. 1983. 
Biological studies of Dall's Porpoise taken 
incidentally by the Japanese salmon mothership 
fishery. International Whaling Commission Scien- 
tific Committee Document SC/35/SM9. 

Jones, L. L., G. C. Bouchet, and B. J. Turnock. 1987. 
Comprehensive report on the incidental take, biology 
and status of Dall's porpoise. International North 
Pacific Fisheries Commission Document No. 3156. 

Kasuya, T. 1978. The life history of Dall's Porpoise 
with special reference to the stock off the Pacific 
coast of Japan. Scientific Reports of the Whales 
Research Institute 30: 1-63. 

Kasuya, T. 1982. Preliminary report of the biology, 
catch and populations of Phocoenoides in the 
western North Pacific. Mammals in the seas, Volume 
4, Food and Agricultural Association Fisheries 
Series 5: 3-19. 

Kasuya, T., and L. L. Jones. 1984. Behavior and 
segregation of the Dall's Porpoise in the northwest- 
ern North Pacific Ocean. Scientific Reports of the 
Whales Research Institute 35: 107-128. 

Kasuya, T., and H. Ogi. 1987. Distribution of mother- 
calf Dall's Porpoise pairs as an indication of calving 
grounds and stock identity. Scientific Reports of the 
Whales Research Institute 38: 125-140. 

Kasuya, T., and S. Shiraga. 1985. Growth of Dall's 
Porpoise in the western North Pacific and suggested 
geographical growth differentiation. Scientific 
Report of the Whales Research Institute 36: 139-152. 

Leatherwood, S., R. R. Reeves, W. F. Perrin, and 
W. E. Evans. 1982. Whales, dolphins, and por- 
poises of the eastern North Pacific and adjacent 
Arctic waters: A guide to their identification. NOAA 
Technical Report NMFS Circular 444: 1-245. 

Miller, E. 1988a. Summary of research on the 
behavior and distribution of Dall's Porpoise, 
Phocoenoides dalli in Puget Sound (May-December, 
1987). Report submitted to the National Marine 
Mammal Laboratory (NOASA, NMFS). 

Miller, E. 1988b. A preliminary application of photo- 
identification techniques to Dall's Porpoise, 



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{Phocoenoides dalli) in Puget Sound. International 
Whaling Commission Scientific Document SC/ A88/ 
P13. 

Morejohn, G. V. 1979. The natural history of Dall's 
Porpoise in the North Pacific Ocean. Pages 45-83 in 
Behaviour of marine animals, Volume 3, Cetaceans. 
Edited by H. E. Winn and B. L. 011a. Plenum Press, 
New York. 

Miyazaki, N. 1987. Biological study on Dall's 
Porpoises incidentally taken by the salmon drift 
gillnet in the land-based fishery area, 1981-1986. 
International North Pacific Fisheries Commission 
Document No. 3146. 

Newby, T. C. 1982. Life history of Dall Porpoise 
{Phocoenoides dalli). True 1885) incidentally taken 
by the Japenese high seas salmon mothership fishery 
in the northwestern North Pacific and western Bering 
Sea, 1978 to 1980. Ph.D. dissertation, University of 
Washington, Seattle. 

Nishiwaki, M. 1967. Distribution and migration of 
marine mammals in the North Pacific area. Bulletin 
of the Ocean Research Institute 17: 93-103. 

Perrin, W. F. 1988. Dolphins, porpoises, and whales. 
An action plan for the conservation of biological 
diversity: 1988-1992. International Union for the 
Conservation of Nature and Natural Resources. 

Pike, G. C, and I. B. MacAskie. 1969. Marine 
mammals of British Columbia. Bulletin of the 
Fisheries Research Board of Canada 171: 1-54. 

Scheffer, V. B. 1949. The Dall's Porpoise, Phoco- 
enoides dalli, in Alaska. Journal of Mammalogy 30: 
116-121. 

Scheffer, V. B. 1950. Porpoises assembling in the 
North Pacific Ocean. Murelet 31: 16. 

Stroud, R. K., C. H. Fiscus, and H. Kajimura. 1981. 
Food of the Pacific white-sided dolphin, Lagenor- 
hynchus obliquidens, Dall's Porpoise, Phocoenoides 



dalli, and northern fur seal, Callorhinus ursinus, off 
California and Washington. Fishery Bulletin (U.S.) 
78:951-959. 

Subramanian, A., S. Tanabe, and R. Tatsukawa. 
1987a. Age and size trends and male-female 
differences of PCBs and DDE in dalli-type Dall's 
Porpoise, Phocoenoides dalli, of northwestern North 
Pacific. Proceedings of the NIPR Symposium on 
Polar Biology 1: 205-216. 

Subramanian, A., S. Tanabe, R. Tatsukawa, S. Saito, 
and N. Miyazaki. 1987b. Reduction in the testoste- 
rone levels by PCBs and DDE in Dall's porpoises of 
the northwestern Pacific. Marine Pollution Bulletin 
18:643-646. 

Tanabe, S., T. Mori, R. Tatsukawa, and N. Miyazaki. 
1983. Global pollution of marine mammals by 
PCBs, DDTs, and HCHs (BHCs). Chemosphere 12: 
1269-1275. 

Tanabe, S., S. Watanabe, H. Kan, and R. Tatsukawa. 
1988. Capacity and mode of PCB metabolism in 
small cetaceans. Marine Mammal Science 4: 
103-124. 

Walker, W. A. 1975. Review of the live-capture 
fishery for smaller cetaceans taken in southern 
California waters for public display, 1966-73. 
Journal of the Fisheries Research Board of Canada 
32: 1197-1211. 

Withrow, D. E., G. C. Bouchet, and L. L. Jones. 1985. 
Response of Dall's Porpoise {Phocoenoides dalli) to 
survey vessels in both offshore and nearshore waters: 
Results of 1984 research. International North Pacific 
Fisheries Commission Document. 

Wursig, B. 1990. Cetaceans and oil: Ecological 
perspectives. In Sea Mammals and Oil: Confronting 
the Risks. Edited by J. R. Geraci, D. J. St. Aubin. 
Academic Press, London (in press). 

Accepted 10 October 1989 



Status of Blainville's Beaked Whale, Mesoplodon densirostrus, 
in Canada* 



J. Houston 



374 Fireside Drive, Woodlawn, Ontario KOA 3M0 

Houston, J. 1990. Status of Blainville's Beaked Whale, Mesoplodon densirostris, in Canada. Canadian 
Field-Naturalist 104(1): 117-120. 

Blainville's Beaked Whale, Mesoplodon densirostris, is widely, if thinly, distributed in tropical and subtropical waters 
and occurs irregularly off the east coast of Canada. It has yet to be reported from the west coast, although one 
stranding has been reported from northern California. The species appears to be more pelagic than other ziphiids and 
of no commercial interest. They are rare in Canadian waters. 

La baleine a bee de Blainville, Mesoplodon densirostris, est une espece cosmopolite, quoique rare, qui frequente les 
eaux tropicales et subtropicales. Elle est parfois presente au large de la cote est du Canada mais elle n'a pas encore ete 
signalee sur la cote ouest quoiqu'on ait observe un individu echoue sur la cote nord de la Californie. L'espece semble 
plus pelagique que les autres Ziphiides et ne porte aucun interet commercial. Elle est rare dans les eaux canadiennes. 

Key Words: Blainville's Beaked Whale, Mesoplodon densirostris. Dense Beaked Whale, baleine a bee de Blainville, 
Cetacea, Ziphiidae, Mesoplodon, status, Canada. 



Blainville's Beaked Whale, Mesoplodon densiros- 
tris (de Blainville 1817), is of medium length 
averaging 4.5 to 5 m and reaching a weight of about 
1000 kg. The species has the spindle shape typical 
of the ziphiids, but can be distinguished by the 
shape of the head (Figure 1). 

The mouth has a distinctive shape formed by the 
two laterally compressed teeth which give the 
mouth a high, arched contour, especially in adult 
whales {see Watson 1981; Leatherwood et al. 
1982). The teeth, in males, may be up to 20 cm in 
height but the root, and much of the crown, are 
enclosed in heavy bone [34% denser than elephant 
ivory (de Blainville 1817)] which gives rise to the 
common name Dense Beaked Whale and the 
specific name densirostris (Watson 1981). This 
feature makes the species easy to distinguish from 
other ziphiids. In addition, the head may be 
flattened in front of the blowhole, providing 
another aid to identification (Leatherwood and 
Reeves 1983). 

The species has small flippers and a small dorsal 
fin situated about midway on the trunk, and it is 
usually pointed and triangular, but may vary in 
shape. There may be a slight convexity on the rear 
edge of the flukes, but a median notch is usually 
absent (Watson 1981; Leatherwood et al. 1982; 
Leatherwood and Reeves 1983). 

Blainville's Beaked Whales are bluish grey to 
dark grey dorsally, being slightly lighter in colour 
ventrally. The anal area may be considerably 
lighter and males in particular have whitish or pink 



blotches over much of the body, thought to be 
caused by "cookie-cutter" sharks or some other type 
of epizoic or parasitic fish (Leatherwood et al. 1982). 
Many males show heavy scarring which may be the 
result of intra-specific fighting {see Heyning 1984). 

Distribution 

Mesoplodon densirostris is the only species of the 
genus which crosses the equator (Davies 1963). The 
species is more widely distributed (Figure 2) than 
any other member of the genus. Blainville's Beaked 
Whales have been reported from the Indian, North 
Atlantic and North and South Pacific Oceans 
(Moore 1966). Although not as yet recorded in the 
South Atlantic, Moore (1966) predicts that it will be 
found there. The species strands more often on 
oceanic islands than on continental coasts, leading 
Moore (1966) to suggest a pelagic distribution, 
further offshore than the other Mesoplodon species. 

The species appears to be relatively uncommon 
along the west coast of North America with only one 
recorded stranding on the northern California coast 
(Leatherwood et al. 1982). On the east coast, 
occasional, single strandings have been recorded 
from Nova Scotia south to Florida, the Bahamas 
and the Gulf of Mexico (Moore 1966; Leatherwood 
and Reeves 1983). 

Protection 

International: Blainville's Beaked Whale is listed 
under Appendix II to the Convention on 
International Trade in Endangered Species 



♦Reviewed and accepted by COSEWIC 11 April 1989 — no designation required. 

117 



118 



The Canadian Field-Naturalist 



Vol. 104 




Figure I. Blainville's Beaked Whale, Mesoplodon densirostris (drawning courtesy of 
Canada Department of Fisheries and Oceans). 



(CITES) of which Canada is a party. Such listing 
requires regulation of trade in live specimens, parts 
and derivatives by those countries party to the 
Convention. Trade must be covered by an 
appropriate export permit issued by the government 
of the exporting country, before entry to another 
party nation is permitted. There are currently over 
100 nations party to the Convention. 

The species is also listed as being Indeterminate in 
Status in the Red Data Book (Goodwin and 
Holloway 1972), and Mitchell (1975a: 13) has 
accepted their listing in placing the species in the 
category of a species not generally known to have 
been taken, or to be presently captured, except for 
scientific purposes, uniquely or accidentally. 

National: This species, along with other members 
of the genus, is protected by general legislation in 
several countries but no specific measures are 
known. 

All cetaceans are protected in Canadian waters 
under the Fisheries Act of 1987 (and amendments 
thereto) and the Cetacean Protection Regulations 
promulgated under the Act. In the United States, 
general protection is provided under the Marine 
Mammals Protection Act of 1972 and the 
Endangered Species Act. 

Population Sizes and Trends 

No reliable information is available on popula- 
tion sizes and trends for this species. Live individuals 
are rarely observed at sea, but have been 
photographed near Hawaii (Leatherwood et al. 
1982). The more pelagic distribution of this species, 
as compared to other members of the genus, may 
indicate that they would be less subject to the coastal 
small whale fisheries which still exist, particularly in 
Japan. The effects of exploitation on numbers may 
therefore be regarded as negligible. Although, 
apparently not abundant where it is known 
(Mitchell i975a), the wide distribution suggests that 
the overall status of the species is secure. 

Habitat 

Little information on habitat preferences, etc. is 
available, as the species is known mainly from 
strandings. These whales appear to be creatures of 



the open seas. Strandings have been recorded more 
often from oceanic islands than on continental 
coasts (Moore 1966; Leatherwood et al. 1982). The 
species appears to have a preference for tropical and 
sub-tropical waters as the distribution seems, for the 
most part, to extend no farther north and/ or south 
of the equator than 35° latitude. The distribution 
may be limited by water temperatures, currents 
and /or distribution of the prey. Their occurrence in 
the North Atlantic as far north as Nova Scotia 
(45° N) may be related to the Gulf Stream (Watson 
1981). 

General Biology 

Nothing is known of the reproductive behaviour 
or life history of the species. The length at birth may 
be between 1.9 to 2.6 m (Leatherwood and Reeves 
1983). Attempts to age individuals reading growth 
layers in the teeth were inconclusive (Mitchell et al. 
1981), but age at maturity appears to be about nine 
years (Leatherwood and Reeves 1983). 

Sound recordings of stranded live animals have 
been made (Poulter 1968; Caldwell and Caldwell 
1971), and members of the species apparently 
produce sounds described as roars, lowing, and 
sobbing and groans. Caldwell and Caldwell (1971) 
described chirps and whistles distinctly pulsed with 
frequencies from 1 to 6 kHz. 

The diet appears to consist of squid and pelagic 
fish (Watson 1981; Leatherwood et al. 1982). 

The species is usually seen in small groups of less 
than 10 individuals (Watson 1981; Leatherwood et 
al. 1982) which may represent family units. 
Blainville's Beaked Whales usually strand singly and 
are often alive when found (Moore 1966; 
Leatherwood et al. 1982). A few sightings have been 
made at sea; the species seems to be shy and has a 
very indistinct blow (Leatherwood et al. 1982). They 
appear to be slow swimmers and dive for periods of 
20 minutes or longer (Watson 1 98 1 ; Leatherwood et 
al. 1982). 

Limiting Factors 

None Known. The species has apparently not 
been subject to exploitation by small whale fisheries, 
and because of its pelagic nature, these are not likely 



1990 



HOUSTON: Status of Blainville's Beaked Whale 



119 



160 MO 120 100 80 60 iO 20 




Figure 2. Distribution of Blainville's Beaked Whale. 



to be a threat in the future. Since small whale 
fisheries are opportunistic and indiscriminate as to 
species, they tend to be multispecies fisheries. In 
such fisheries one may think that a falling catch per 
unit effort would not influence future catch effort. 
However, many of the species in such fisheries are 
rare and it cannot be assumed that any future 
exploitation would, in the longterm, be inconse- 
quential (see Mitchell 1975b). At present, the species 
is probably inherently protected by its rarity and 
pelagic habits. 

Special Significance of the Species 

Most of what is known of ziphiids in Canadian 
waters comes from the few specimens stranded here. 
Blainville's Beaked Whale has not as yet been 
recorded from British Columbia but is known from 
Nova Scotia. The species is not known to have been 
taken by commercial whalers and probably not by 
coastal fisheries because of its pelagic habits. It is 
doubtful that the species or parts or derivatives 
would show up in international trade as there is no 
demand. 

Blainville's Beaked Whale could be confused with 
Cuvier's Beaked Whale {Ziphius cavirostris) where 
the ranges overlap. Identification requires an expert. 

Evaluation 

Blainville's Beaked Whales are medium-sized 
cetaceans which are widely dispersed in tropical and 
sub-tropical waters. Mitchell (1975a, b) has 
indicated that most species of the genus are probably 
not abundant and that even moderate exploitation 
could impact negatively on existing populations. 
The species is not known from British Columbia and 
is only rarely seen on the east coast. 



Given the above, and the fact that Canadian 
distribution is on the periphery of the range, the 
species would appear to be rare in Canadian waters. 
As it is not under any threat in Canada there is no 
reason to consider a COSEWIC status designation 
for the species at this time. 

Acknowledgments 

Production of this report was made possible 
through the support of the Department of Fisheries 
and Oceans, the Canadian Wildlife Service and 
World Wildlife Fund Canada. 

Literature Cited 

Caldwell, D. C, and M. C. Caldwell. 1971. Sounds 
produced by two rare cetaceans studied in Florida. 
Cetology 4: 1-6. 

Davies, J. L. 1963. The antitropical factor in cetacean 
speciation. Evolution 17: 107-116. 

de Blainville, M. 1817. Nouveau Dictionaire d'Histoire 
Naturelle, Paris, 9: 178-179. 

Goodwin, H. A., and C. W. Holloway. 1972. Red data 
book, Volume I (Mammaha). International Union for 
Conservation of Nature and Natural Resources, 
Morges, Switzerland. 

Heyning, J. E. 1984. Functional morphology involved in 
intraspecific fighting of the beaked whale Mesoplodon 
carlhubbsi. Canadian Journal of Zoology 62: 
1645-1654. 

Leatherwood, S., and R. R. Reeves. 1983. The Sierra 
Club handbook of whales and dolphins. Sierra Club, 
San Francisco, California. 

Leatherwood, S., R. R. Reeves, W. F. Perrin, and W. E. 
Evans. 1982. Whales, dolphins and porpoises of the 
eastern North Pacific and adjacent Arctic waters. U.S. 
Department of Commerce National Oceanic and 
Atmospheric Administration, National Marine 
Fisheries Service Technical Report Circular 444. 



120 



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Vol. 104 



Mitchell, E. 1975b. Porpoise, dolphin and small whale 
fisheries of the world. International Union for 
Conservation of Nature and Natural Resources, 
Morges, Switzerland, Monograph Number 3. 

Mitchell, E. 1975b. Report of the Meeting on Smaller 
Cetaceans, Montreal, April 1-11, 1974, Subcommittee 
on Small Cetaceans, Scientific Committee, Interna- 
tional Whaling Commission. Pages 889-984 in Review 
of biology and fisheries for smaller cetaceans. Edited by 
E. Mitchell. Journal of the Fisheries Research Board of 
Canada 32(7): Special Issue. 

Mitchell, E. D., J. G. Mead, and V. M. Kozicki. 1984. 
Abstract. Readability of growth layers in teeth of beaked 
whales, Ziphiidae. Reports of the International Whaling 
Commission, Special Issue Number 3: 215. 



Moore, J. C. 1966. Diagnoses and distributions of 
beaked whales of the genus Mesoplodon known from 
North American waters. Pages 31-61 in Whales, 
dolphins and porpoises. Edited by K. S. Norris. 
University of California Press, Los Angeles, 
California. 

Poulter, T. C. 1968. Marine mammals. Pages 405-465 
in Animal communication; techniques of study and 
results of research. Edited by T. A. Sebeok. Indiana 
University Press, Bloomington, Indiana. 

Watson, L. 1981. A sea guide to whales of the world. 
Nelson Canada Limited, Scarborough, Ontario. 



Accepted 10 October 1989 



Status of Hubbs' Beaked Whale, Mesoplodon carlhubbsi, 
in Canada* 



J. Houston 

374 Fireside Drive, Woodlawn, Ontario KOA 3M0 

Houston, J. 1990. Status of Hubbs' Beaked Whale, Mesoplodon carlhubbsi, in Canada. Canadian Field-Naturalist 
104(1): 121-124. 

Hubbs' Beaked Whale, Mesoplodon carlhubbsi, is a rare member of the North Pacific fauna. The species is known 
from 3 1 stranded specimens and one possible live sighting. Most strandings have been along the North American coast 
from Prince Rupert, British Columbia to La Jolla, California. Four strandings are recorded from Ayukawa, Japan. 
The species is not known to have been, or to be, of interest to commercial fisheries and is probably protected by its 
rarity and occurrence in less frequented (by man) waters of the North Pacific. 

La baleine a bee de Hubbs, Mesoplodon carlhubbsi, est une espece rare de la faune du Pacifique Nord. On connait 
I'espece d'apres 3 1 specimens echoues et une observation possible d'un specimen vivant. La plupart des echouements se 
sont produits le long de la cote de I'Amerique du Nord, depuis Prince Rupert, en Colombie-Britannique, jusqu'a La 
Jolla, en California. On signale par ailleurs quatre specimens echoues a Ayukawa, au Japon. L'espece n'est pas 
reconnue pour avoir presente ou pour presenter un interet pour la peche commerciale et est probablement protegee en 
raison de sa rarete et de sa presence dans les eaux moins frequentees (par les hommes) du Pacifique Nord. 

Key Words: Hubbs' Beaked Whale, Mesoplodon carlhubbsi, Arch-beaked Whale, baleine a bee de Hubbs, Cetacea, 
odontocetes, Ziphiidae, beaked whales, Mesoplodon. 



Hubbs' Beaked whale, Mesoplodon carlhubbsi 
Moore 1963, is a toothed whale of the family 
Ziphiidae and is typical of the genus in size and 
shape (see Watson 1981). Also known as the Arch- 
beaked Whale, this small ziphiid was not described 
as a separate species until the 1960s (Moore 1963). 
The species is similar enough to Mesoplodon 
stejnegeri (Stejneger's Beaked Whale) to be 
confused with it. It is known mainly from stranded 
specimens (Mead et al. 1982). 

These are medium-sized beaked whales (Figure 
1) with ipaximum length estimated at about 5.3 m 
and a maximum weight of about 1500 kg (Watson 
1981; Meadetal. 1982). The shape is similar to that 
of other ziphiids with a large thorax and small head 
and tail. There is a prominent pair of vertical 
throat grooves; the flippers are thin and elongated 
and can be depressed in "flipper pocket" 
repressions in the body wall posterior to each 
flipper. There is no median notch on the flukes 
which are otherwise similar to those of most 
medium-sized whales (Watson 1981; Mead et al. 
1982). 

The most striking external features of the species 
is the pigmentation of the head, particularly in 
adult males. The white rostrum contrasts with the 
nearly black body, and on the mandible, the 
posterior edge of the white patch parallels the 
posterior edge of the erupted tooth (Mead et al. 



1982; Heyning 1984). There is less contrast in 
females and subadults but the area is generally 
lighter than the remainder of the head. The extent 
of the lighter pigmentation may be age related 
(Mead et al. 1982). The rest of the body is dark grey 
to black but the ventral surface may be lighter in 
subadults and females. The bodies of adult males 
tend to be marked by uneven linear scars which are 
thought to result from intra-specific fighting 
(Heyning 1984). As in all Mesoplodon species, the 
teeth erupt only in adult males and the rostum is 
strongly constricted by the projecting pair of 
mandibular teeth (Mead et al. 1982) which are 
larger than in other ziphiids (Heyning 1984). 

Distribution 

Most of the information concerning distribution 
comes from the relatively few (31) recorded 
strandings {see Mead et al. 1982). The species 
seems to be restricted to the North Pacific (Figure 
2) and most strandings (27) have been along the 
west coast of North America from San Diego, 
Califoria (33° N) to Prince Rupert, British 
Columbia (54° N). This may be the northern limit 
as no strandings have been reported north of 54°N, 
but there are abundant records of Mesoplodon 
stejnegeri north of this limit. The southern limit 
may be the product of a general lack of 
information, as no strandings have been recorded 



♦Reviewed and accepted by COSEWIC 11 April 1989 — no designation required. 



121 



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Figure I. Hubbs' Beaked Whale, Mesoplodon carlhubbsi {redrsL-wn and adapted from 
Watson 1981). 



from Central America south to the equator (Mead 
et al. 1982). Pike and Mac Askie (1962) report three 
strandings along the British Columbia coast and 
Mead et al. (1982) recorded four others. One 
sighting of a small whale by Hubbs (1946) off La 
Jolla, California, is the only posible sighting of a 
live animal. 

The range appears to extend west to Japan 
where the species has been recorded from the 
northeast coast of Honshu at about 38° N latitude 
(Mead et al. 1982). The range appears to be south 
of that of Stejneger's Beaked Whale and north of 
that of Mesoplodon ginkgodens, the Ginkgo- 
Toothed Beaked Whale (Watson 1981). 

Protection 

International: Mesoplodon species are included 
in Appendix II of the Convention on International 
Trade in Endangered Species (CITES). No other 
international agreements refer to the genus. The 
species is listed as 'Indeterminate' in the Red Data 
Book (Goodwin and Holloway 1972). 

National: The genus is protected by general 
legislation in several countries but no specific provi- 
sions are known. In Canada, general protection is 
accorded under the Fisheries Act and the Cetacean 
Protection Regulations which prohibit commercial 
whaling. In the United States, general protection is 
accorded under the Marine Mammal Protection 
Act of 1972 and the Endangered Species Act. 

Population Sizes and Trends 

Known only from 31 stranded specimens and one 
possible live sighting, this species can only be 
described as rare. The species is not known to have 
been commercially exploited or captured (Mitchell 
1975). No other information is available on 
population status. 



Habitat 

The distribution of Mesoplodon carlhubbsi along 
the Japanese coast appears to coincide with the 
surface Transition Domain and at depth with the 
Subarctic Current System where deep elements of 
the Kuroshio and Oyashio currents mix. The North 
American distribution coincides with the surface 
Dilute and Upwelling domains and with the 
confluence of the Subarctic and California current 
systems at depth (Favorite et al. 1976; Mead et al. 
1982). 

The distribution is probably more directly related 
to the distribution of the prey species on which it 
feeds (mesopelagic squid and fish) than to the 
character of the water mass (Mead et al. 1982). 
However, the distribution of the prey may be 
directly related to the character of the intermediate 
and deep water masses. 

General Biology 

Although only a limited number of specimens 
have been examined, length at physical maturity 
appears to be at around 5 m. Presumably, growth 
occurs more slowly after physical maturity, until a 
maximum observed length of 5.3 to 5.4 m is reached 
(Mead et al. 1982; Mead 1984). To date, there are no 
estimates of longevity. 

Examination of neonate specimens (from 
strandings) indicates that calving probably takes 
place in the summer and that the gestation period is 
about twelve months, as for other small cetaceans 
(Mead et al. 1982). No information is available on 
breeding or calving and it is not known if the species 
makes seasonal migrations. The largest recorded 
fetus measured 0.9 m and the smallest calf 2.47 m; 
the mean length at birth has been estimated to be 2.5 
m(Mead 1984). 



1990 



HOUSTON: Status of Hubbs' Beaked Whale 



123 



160 140 120 100 aO 60 40 20 




Figure 2. Distribution of Hubbs' Beaked Whale. 



The linear scars on the bodies of adult males have 
been attributed to aggressive use of the teeth by 
other males of the same species (Heyning 1984). 
Heyning suggests that the structure of the teeth has 
evolved in relation to social interactions involving 
the establishment of breeding territories rather than 
for the acquisition and manipulation of food items. 
Stomach contents retrieved from stranded 
specimens suggest a diet of squid and fish (Mead et 
al. 1982). 

No information is available on internal parasites 
or diseases. Externally, the species may be 
parasitized by the copepod Penella (Ivashin and 
Golubovsky 1978) and possibly balanomorph and 
lepadomorph barnacles (Mead et al. 1982). 

Limiting Factors 

None known. Rarity is the present main 
protection of the species, in concert with a 
distribution in less frequented waters and 
unexploited food species. 

Special Significance of the Species 

Hubbs' Beaked Whale has only recently been 
described (Moore 1963) and can easily be confused 
with other members of the genus, particularly 
Stejneger's Beaked Whale and Andrew's Beaked 
Whale {Mesoplodon bowdoini) where the ranges 
overlap (Watson 1981). The species is not known to 
have been hunted, although occasional specimens 
might be taken during other fishing operations. 
None has been reported in captivity, but study of 
captive individuals, if possible, would provide useful 
information, otherwise difficult to obtain in the 
wild. 

The only parts and derivatives that would be 
likely to appear in trade would be scientific 



specimens. It is unlikely that such a rare species 
could support a fishery or would be of commercial 
interest. 

Evaluation 

The species is a member of the pelagic fauna of the 
Pacific waters of Canada. Although nothing is 
known of its habit or life history in Canadian waters, 
it appears to be a rare component of this ecosystem. 
The species is under no imminent threat in Canadian 
waters and COSEWIC status is probably neither 
warranted nor required at this time. 

Acknowledgments 

I am indebted to the World Wildlife Fund 
(Canada), the Department of Fisheries and Oceans, 
and the Canadian Wildlife Service, for financial 
support in the production of this report. Thanks are 
also extended to COSEWIC for the opportunity to 
present the information and for the support and 
encouragement of the Fish and Marine Mammals 
Subcommittee. 

Literature Cited 

Favorite, F., A. J. Dodimeade, and K. Nasu. 1976. 

Oceanography of the Subarctic Pacific Region. 

International North Pacific Fisheries Commission 

Bulletin 33. 
Goodwin, H. A., and C. W. Holioway. 1972. Red Data 

Book, Volume I (Mammaha). International Union for 

the Conservation of Nature, Morges, Switzerland. 
Heyning, J. E. 1984. Functional morphology involved in 

intraspecific fighting of the beaked whale, Mesoplodon 

carlhubbsi. Canadian Journal of Zoology 62: 1645- 

1654. 
Hubbs, C. L. 1946. First records of two beaked whales, 

Mosoplodon bowdoini and Ziphius cavirostris, from 

the Pacific coast of the United States. Journal of 

Mammalogy 27: 247-255. 



124 



The Canadian Field-Naturalist 



Vol. 104 



Ivashin, M. V., and Yu. P. Golubovsky. 1978. On the 
cause and appearance of white scars on the body of 
whales. Reports of the International Whaling 
Commission 28: 199. 

Mead, J. G. 1984. Survey of reproductive data for the 
beaked whales (Ziphiidae). Reports of the International 
Whaling Commission, Special Issue 6: 91-96. 

Mead, J. G., W. A. Walker, and W.J. Houck. 
1982. Biological observations on Mesoplodon carlhub- 
bsi (Cetacean: Ziphiidae). Smithsonian Contributions 
to Zoology Number 344. 

Mitchell, E. 1975. Porpoise, dolphin and small whale 
fisheries of the world: status and problems. Inter- 



national Union for the Conservation of Nature, 

Monograph Number 3. 
Moore, J. C. 1963. Recognizing certain species of beaked 

whales in the Pacific Ocean. American Midland 

Naturalist 70: 396-428. 
Pike, G. C, and L B. MacAskie. 1969. Marine mammals 

of British Columbia. Fisheries Research Board of 

Canada Bulletin 171. 
Watson, L. 1981. Sea guide to whales of the world. 

Nelson Canada Limited, Scarborough, Ontario. 



Accepted 10 October 1989 



Status of Sowerby's Beaked Whale, Mesoplodon bidens, 
in Canada* 



Jon Lien and Frances Barry 



Department of Psychology and Newfoundland Institute for Cold Ocean Science, Memorial University of 
Newfoundland, St. John's, Newfoundland AlB 3X9 

Lien, Jon, and Frances Barry. 1990. Status of Sowerby's Beaked Whale, Mesoplodon bidens, in Canada. Canadian 
Field-Naturalist 104(1): 125-130. 

Sowerby's Beaked Whale of North Sea Beaked Whale, Mesoplodon bidens, has been encountered only 1 1 times in the 
western North Atlantic through two mass strandings, seven strandings of individuals and in two sightings. There are 
more recorded strandings of this whale on British and European coasts but its range appears to be generally offshore 
throughout the North Atlantic. Because of the paucity of encounters with Mesoplodon bidens, little is known of its 
biology. 

La baleine a bee de Sowerby (dauphin du Havre), Mesoplodon bidens, n'a ete repere que onze fois dans I'ouest de 
I'Atlantique Nord, lors de deux echouages en masse, sept echouages d'individus isoles et deux observations. Bien que 
les releves d'echouages soient plus frequents sur les cotes britanniques et europeennes, I'aire de repartition de cette 
espece semble couvrir generalement en haute mer tout I'Atlantique Nord. La biologic de Mesoplodon bidens est peu 
connue en raison du nombre extremement reduit d'observations. 

Key Words: Sowerby's Beaked Whale, Mesoplodon bidens. North Sea Beaked Whale, dauphin du Havre, baleine a 
bee de Sowerby, Cetacea, odontocetes, ziphiids, beaked whales, Mesoplodon, rare and endangered species. 



Mesoplodon bidens (Sowerby 1804), Sowerby's 
Beaked Whale, is also known as the North Sea 
Beaked Whale. The former name refers to the 
describer of the first specimen found in Scotland in 
1800 (Katona et al. 1983). Little is known of its 
distribution and biology. Only a few papers discuss 
the species, usually incidentally, in general reports 
of stranded cetaceans. 

Early reviews of the beaked whales in general 
(Flower 1872, 1878; True 1910) were based on 
limited numbers of specimens and several species 
were not recognized. The genus Mesoplodon 
comprises twelve different species, most of which 
are poorly known and generally regarded as rare 
(Leatherwood and Reeves 1983). Seven species are 
now recognized in the Northern Hemisphere 
(Moore 1966) and occur in both Atlantic (Ulmer 
1941) and Pacific waters (Orr 1953). However, two 
species are found only in the Pacific, Mesoplodon 
stejnegeri and Mesoplodon bowdoini (Moore 
1966). Occurrence of Mesoplodon bidens is 
restricted to the North Atlantic. General reviews of 
beaked whales of the genus Mesoplodon are given 
by Moore (1966) and Leatherwood and Reeves 
(1983). 

Mesoplodon bidens is a medium-sized beaked 
whale known to reach 5 m in length (Figure 1). 
Adults are dark charcoal grey on the back with 
light spots overall. Young animals are also dark 
charcoal grey on the back but are lighter on the 



belly and unspotted (Leatherwood et al. 1976). The 
unnotched flukes of adults are dark above and 
below. The spindle shaped body has relatively long 
pectoral flippers which are about 1/8 body length. 
Family characteristics include the long narrow 
snout and v-shaped slashes or grooves on the 
throat (Figure 2). Positive field identification of 
most species is difficult, but may be simplified by 
comparisons of such characteristics as mandible 
size and tooth position where ranges overlap (see 
Fraser 1953 or Watson 1981). Each side of the 
lower jaw has one triangular tooth. Teeth erupt 
above the gum in males but not in females. Positive 
diagnosis of stranded specimens can be based on 
skull characteristics (Moore 1966). 

Distribution 

The range of Mesoplodon bidens, the most 
northerly species of beaked whale, is poorly known 
but distribution estimates based on stranding 
events and a few sightings have been made. 

There have been 30 or so strandings of this 
species on the European coast (Saemundsson 
1939; Fraser 1953) and occasional sightings (^ynes 
1974; Christensen 1977). Saemundsson (1939) and 
Christensen (L Christensen, Institute of Marine 
Research, Bergen-Nordnes, Norway; personal 
communication) indicate occasional captures by 
Norwegian whalers off Iceland and in the Barents 
Sea. Sheldrick (1979) reports 29 stranding records 



* Vulnerable status approved and assigned by COSEWIC 11 April 1989. 

125 



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The Canadian Field-Naturalist 



Vol. 104 




Figure 1. Drawing of an adult male Sowerby's Beaked Whale, Mesoplodon bidens (redrawn from 
Watson 198 1). 



of Mesoplodon bidens on the British coastUne 
from 1913 to 1972. Moore (1966) concluded that 
the relative abundance of Mesoplodon bidens 
strandings on the European side of the Atlantic, in 
a gross way, was related to the greater abundance 
of animals living in that vicinity compared to the 
Western Atlantic. Saemundsson (1939) suggested 
the species has an open sea distribution with 
occasional visits to coastal waters. 

Recent records from eastern Canada (Sergeant 
and Fisher 1957; Dix et al. 1986; Lien et al. 1990) 
suggest that the animals frequent waters off this 
coast fairly regularly. Altogether, seven individual 
strandings, two mass strandings involving a total 
of nine whales, and two live sightings of 
Mesoplodon bidens have been recorded from 1906 
to 1988 in the Western North Atlantic. These are 
listed in Table 1 and locations are shown in Figure 
3. Most of these events have been recorded in the 
last 15 years and over half in the past four years. 
This does not likely indicate an influx or 
redistribution of the species to the western North 
Atlantic, but rather reflects recent increases in the 
efficiency of North American stranding networks 
(Lien et al. 1990) as well as verifiable field 
identification through the use of photography. 

Mesoplodon bidens was first recorded in the 
Northwest Atlantic on U.S. shores in 1867 when an 
adult male was stranded on Nantucket Island, 
Massachusetts (Allen 1906). A stranding of a 488 
cm female occurred at Nantucket, Massachusetts 
in September 1982. The most recent U.S. stranding 
was of a 457 cm male which occurred at Port St. 
Joe on the Gulf Coast of Florida in October 1984 
(J. G. Mead, Division of Mammals, Smithsonian 
Institution, Washington, D.C.; personal com- 
munication). 

The remaining specimens of Mesoplodon bidens 
in North America were all recorded in Newfound- 
land and Labrador. Two were discovered in 
consecutive summers (Sergeant and Fisher 1957); a 
472 cm adult male was found dead in August 1952 
in Trinity Bay (47°45'N, 53°52'W), and a 427 cm 
immature female was harpooned in Notre Dame 
Bay (49°40'N, 55°50'W) in September 1953. A 
third partial animal, probably a female, came from 
Labrador (54°I0'N, 58°35'W) in September 1973 



(J. G. Mead, personal communication). A 410 cm 
male was found dead in July 1984 in Conception 
Bay, Newfoundland (47°34'N, 53°11'W) and was 
assumed (from photographs) to be the same whale 
which had been sighted and released from fishing 
gear two days previously (Dix et al. 1986). 

Mass strandings of Mesoplodon bidens have 
occurred for two consecutive years on the 
northeast coast of Newfoundland. Three whales, 
all males 462, 485, and 495 cm in length, died and 
were examined from a group of six individuals 
which stranded in August 1986 near Carmanville, 
Newfoundland (49°07'N, 54°18'W) [Lien et al. 
1990]. In September 1987, reports of three whales 
beaching on several occasions near Norris Arm, 
Newfoundland (49°07'N, 55°15'W) led to the 
discovery and examination of a single 462 cm 
female (Lien et al. 1990). 

Protection 

Mesoplodon bidens is listed in Appendix I of the 
Convention on International Trade in Endangered 
Species of Wild Flora and Fauna (CITES). In 
waters under Canadian jurisdiction, Mesoplodon 
bidens is protected by the Fisheries Act. In the 
United States, it is classified as endangered and 
protected under the Marine Mammal Protection 
Act. 

Population Size and Trends 

There are no estimates of the population size of 
Mesoplodon bidens due to insufficient data. The 
very scantiness of data on the species probably 
indicates this whale is quite rare. 

Following the inception of the comprehensive 
stranding network in Newfoundland in 1979 (Lien 
1980; Lynch 1988), verified sightings or strandings 
of Mesoplodon bidens individuals or groups have 
averaged 0.30 per year. This compares to an 
average of 0.12 sightings or strandings per year 
during the period 1947-1973 (Sergeant and Fisher 
1957; Sergeant et al. 1970; Dix et al. 1986). Prior to 
this early work there were no encounters recorded 
with ziphiids. 

Stranding networks have become more efficient 
in many areas of North America recently (Geraci 
and St. Aubin 1979; Lien 1980), and because of the 



1990 



Lien and Barry: Status of Sowerby's Beaked Whale 



127 




Figure 2. Photograph oi the 362 cm female stranded at 
Bay of Exploits, Newfoundland in September 
1987 (see Table 1). Note the "V" shaped throat 
groove. 

presence of the unusual rostrum, it is likely that 
most stranded Mesoplodon bidens would be 
reported. Although the number of strandings 
reported in recent years has increased, it can not be 
concluded that Mesoplodon bidens has become 
more common in Canadian waters in the past 
decade. The increased scope and efficiency of 
present stranding networks, where effort is 
difficult to quantify, may well bias conclusions 
about the relative abundance of the species. 



However, earUer suggestions that the distribution 
of Mesoplodon bidens enters in the North Sea and 
only occasionally wanders to western North 
Atlantic waters (Moore 1966) should perhaps be 
regarded as premature. 

Habitat 

There are, at present, inadequate data to identify 
habitat requirements of Mesoplodon bidens. 

Sightings of ziphiids have been common along 
the edges of temperature fronts in water depths of 
approximately 200 m (Price and Fairfield 1985) 
which is consistent with a sighting of Mesoplodon 
bidens by Marion et al. (1987). However, 
Mesoplodon bidens sightings have also occurred in 
waters of depths greater than 1500 m ((/)ynes 1974; 
Christensen 1977). 

If Mesoplodon bidens is a pelagic species that 
only occasionally visits coastal waters, one might 
expect mass strandings of the species (Sergeant 
1982; Tyack 1987). In the past three decades, the 
ratio of mass to single Canadian strandings of 
Mesoplodon bidens is 2: 1 . For British shores, the 
mass to single stranding ratio of Mesoplodon 
bidens is 1:29 (Sheldrick 1979). Sheldrick (1979) 
makes the point that mass strandings are 
generally considered rare for all species on British 
coasts. 



Table 1. Records of Me.$o/7/o(io« fe/Wen^ in North America (1867-1' 



Date 


Location 


Details 


Source 


1867 


Nantucket Island, Massachusetts 


male 


Allen (1906) 


25 Aug. 1952 


Chapel Arm, Trinity Bay, 


472 cm male 


Sergeant and 




Newfoundland 


NMC-26483 


Fisher (1957) 




(47°45'N, 53°52'W) 






23 Sept. 1953 


Wild Bight, Notre Dame Bay, 


472 cm female 


Sergeant and 




Newfoundland 


NMC-26484 


Fisher 1957 




(49°40'N, 55°50'W) 






Sept. 1973 


Labrador, Notre Dame Bay, 


part of 


J. Mead (personal 




(54°10'N, 58°35'W) 


female 


communication) 


10 Sept. 1973 


Nantucket, Masschusetts 


488 cm male 


J. Mead (personal 






MH-82-I80 


communication) 


24 July 1984 


Manuels, Conception Bay, 


net entrapment 


Dix et al. 




Newfoundland 




(1986) 




(47°35'N, 53°11'W) 






26 July 1984 


Port de grave. Conception Bay, 


410 cm male 


Dix et al. 




Newfoundland 




(1986) 




(47°34'N, 53°11'W) 






Oct. 1984 


Port St. Joe, Gulf coast of Florida 


457 cm female 


J. Mead (personal 






USNM 550414 


communication) 


30 Aug. 1986 


Carmenville, Newfoundland 


mass stranding of 6 


Lien et al. 




(49°07'N, 54°I8'W) 


whales; 3 males 495 cm, 
485 cm, 495 cm. (Cana- 
dian Museum of 
Nature) 


(1989) 


July 18 1987 


Hydrographers Canyon region 


sighting — 3 adults and 


Marion et al. 




(40°00'N, 68°54'W) 


2 calves 


(1987) 


Sept. 18 1987 


Norris Arm, Bay of Exploits, 


mass stranding of 3 


Lien etal. (1989): See 




Newfoundland 


whales; one 362 cm 


Figures 2,3 




(49°07', 55°15'W) 


female, Ontario Science 
Centre 





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The Canadian Field-Naturalist 



Vol. 104 



90 


75 

Canada 


/ 




f 


^^^^ 




y 


^ 


u^ 


45 - 


L 

United States 


J 




Atlantic 




of America 


\ 




Ocean 


30- 


"■".:.. V 


W 









Figure 3. Locations of North American sightings and 
strandings of Mesoplodon bidens, 1867-1988. 



Another species, the Long-finned Pilot Whale 
{Globicephala melaena), which is an offshore 
pelagic species and returns to coastal Canadian 
waters seasonally (Mercer 1975), may be somewhat 
comparable. For the period 1957 to 1980, Sergeant 
(1982) lists 12 mass strandings of Globicephala 
melaena in eastern Canada which provide a mass to 
single strandings ratio of about 1:1. The high ratio of 
mass to single strandings in Newfoundland (Lien et 
al. 1990) could be interpreted as support for the 
hypothesis that Mesoplodon bidens is an offshore 
species which tends to mass-strand rather than to 
strand individually, and that waters off Newfound- 
land are in the centre of the species range rather than 
at its edge (Sergeant 1982). 

Sergeant and Fisher (1957) suggest that, 
following the habit of the Long-finned Pilot 
Whale, Mesoplodon bidens occurs inshore when 
the squid Illex illecebrosus is abundant. However, 
Dix et al. (1986) note that strandings of 
Mesoplodon bidens occur in years of low squid 
abundance and no trace of squid has been 
discovered in stomachs of specimens examined. 
Moreover, examination of oceanographic data 
from the years of strandings has revealed no 
obvious correlations with, or differences from 
other years which might account for the inshore 
appearance of Mesoplodon bidens (Dix et al. 
1986). 

Biology 

Very little is known about the life history of 
Mesoplodon bidens. The limited knowledge 
available has been derived from stranded 
specimens and the few verified field sightings. 



Age of breeding is unknown. Females with 
overall lengths of 483 cm (Reiner 1986), and 505 
cm (Jonsgard and Hoidal 1957) were mature, 
whereas one 462 cm female was immature (Lien et 
al. 1990). Males under 5 m have been considered 
immature (Sergeant and Fisher 1957; Dix et al. 
1986; Lien et al. 1990). Mead (1984) reports a 
maximum recorded length of 5.5 m for males. 
Fetuses of 54 cm (Reiner 1986), 158, and 121 cm 
have been reported, and Greig (1908) found a very 
young calf measuring 245 cm in length with a 
remnant of umbilical cord (Greig 1908). Jonsgard 
and Hoidal (1957) conclude that calves must be 
from 150 to 250 cm in length, most likely between 
180 to 210 cm at birth and that mating and birth 
occur in late winter and early spring after a 
gestation of about one year. They also suggest, 
based on what was assumed to be a mother and 
calf, that weaning may occur after about a year 
when the calf is about 300 cm in length. 

There are too few observations of Mesoplodon 
bidens to infer much about its movements and 
behaviour. 

Ziphiids are usually found singly, or in small 
groups and Mesoplodon species, in particular, 
have been found to have the least social cohesion of 
the medium-sized odontocetes (Gaskin 1982). 
Some observations of Mesoplodon bidens appear 
to support this view. 

Groups of Mesoplodon bidens may be segre- 
gated by sex. The low probability of a mass 
stranding of three males from a group of six 
individuals suggests the hypothesis that all-male 
social groups occur (Lien et al. 1990). The 
possibility of sexual segregation in Mesoplodon 
bidens raises hypotheses explaining their social 
structure. Gaskin (1982) has related segregation by 
sexes to differences in food requirements of males 
and females or to their polygynous mating habits. 
However, comparisons of body measurements 
from a limited number of males and females 
indicates little sexual dimorphism in Mesoplodon 
bidens (Lien et al. 1990), except in the degree of 
development of the single pair of mandibular teeth 
(Moore 1966). The teeth erupt only in males, while 
females have smaller teeth which do not protrude 
from the gums (Sergeant and Fisher 1957). This 
minimal degree of sexual dimorphism suggests 
that polygyny may not be the mating organization 
(Davies and Kreb 1981; Sergeant 1982). 

Limiting Factors 

Unknown. 

Special Significance of the Species 

Mesoplodon bidens has only been reported 1 1 
times in North American waters in the past 120 
years. Sightings and strandings are rare compared 



1990 



Lien and Barry: Status of Sowerby'S Beaked Whale 



129 



to those of other ziphiids and cetaceans generally. 
Study of the western North Atlantic population is 
very important to help understand this species and 
virtually any information about these whales will 
greatly increase our knowledge about their lives. 

Evaluation 

There is insufficient evidence to estimate 
population size and range. Since information on 
food and habitat requirements is also speculative, 
trends of populations of this species are impossible 
to discern or predict. Due to lack of knowledge of 
Mesoplodon bidens, and given the rarity of 
encounters, this species should be considered rare 
throughout its range until more information is 
available. Due to its rarity and limited Canadian 
distribution coincidental with major shipping 
lanes and areas of offshore petrochemical 
exploration and development, the species should 
be considered vulnerable in Canadian waters. 

Acknowledgments 

Support for this work was made available 
through the Department of Fisheries and Oceans 
and World Wildlife Fund (Canada). We thank 
Wayne Ledwell and Jennifer Nauen for their 
assistance in completing this manuscript and 
Dawn Nelson for her drawing. 

Literature Cited 

Allen, G. M. 1906. Sowerby's whale on the American 

coast. American Naturalist 40: 357-367. 
Christensen, I. 1977. Observations of whales in the 

North Atlantic. Report of the International Whaling 

Commission 27: 388-399. 
Davies, N. B., and J. R. Krebs. 1981. Ecology, natural 

selection and social behaviour. Pages 1-18 in 

Behavioural ecology. An evolutional approach. Edited 

by J. R. Krebs and N. B. Davies. Sinauer Associates, 

Inc. 
Dix, L., J. Lien, and D. E. Sergeant. 1986. A North Sea 

Beaked Whale, Mesoplodon bidens, in Conception 

Bay, Newfoundland. Canadian Field-Naturalist 

100(3): 389 391. 
Flower, W. H. 1872. On the recent ziphioid whales with 

a description of the skeleton of Berardius arnouxi. 

Transactions of the Zoological Society, London 8: 

203-234. 
Flower, W. H. 1878. A further contribution to the 

knowledge of the ziphioid whales, genus Mesoplodon. 

Transactions fo the Zoological Society, London 10: 

415-437. 
Fraser, F. C. 1953. Report of Cetacea stranded on the 

British coasts from 1938-1947. British Museum 

(Natural History) Number 13: 38-40. 
Gaskin, D. E. 1982. The ecology of whales and 

dolphins. Heinemann, London. [Pages 1 19, 123, 135.] 
Geraci, J. R., and D.J. St. Aubin. 1979. Biology of 

marine mammals: insights through strandings. U.S. 

Marine Mammal Commission, Washington, D.C. 343 

pages. 



Greig, J. A. 1908. Spidshvalen. Norsk Fiskeri Tidende 
6te hefte: 264-268. 

Jonsgard, A., and P. Hoidal. 1957. Strandings of 
Sowerby's Beaked whale {Mesoplodon bidens) on the 
west coast of Norway. Norsk Hvalfangst-Tidende 9: 
507-512. 

Katona, S. K., V. Rough, and D. Richardson. 1983. A 
field guide to the whales. Porpoises and seals of the 
Gulf of Maine and eastern Canada: Cape Cod to 
Newfoundland. Third Edition. Scribner, New York. 
255 pages. 

Leatherwood, S. D., and R. R. Reeves. 1983. The 
Sierra Club handbook of whales and dolphins. Sierra 
Club Books, San Francisco, California. 302 pages. 

Leatherwood, S., D. Cadwell, and H. E. 
Winn. 1976. Whales, dolphins and porpoises of the 
western North Atlantic: a guide to their identification. 
U.S. Department of Commerce, National Oceanic and 
Atmospheric Administration, National Marine 
Fisheries Service. 176 pages. 

Lien, J. 1980. Ma^w^cn/?;. Whale collisions with fishing 
gear in Newfoundland. Final report to Fisheries and 
Oceans Canada, Ottawa, Ontario. 316 pages. 

Lien, J., F. Barry, K. Breeck, and U. Zusch- 
lag. 1990. Mass strandings of Sowerby's Beaked 
Whales {Mesoplodon bidens) in Newfoundland. 
Canadian Field-Naturalist 104(3): in press. 

Lynch, K. D. 1988. Humpback, finback, minke and 
pilot whale distributions in Newfoundland and 
Labrador 1976-1983. M.Sc. thesis. Memorial 
University of Newfoundland, St. John's, Newfound- 
land. 196 pages. 

Marion, S., S. Forhock, and P. Fontaine. 1987. First 
sighting of Mesoplodon bidens in North American 
waters. North Atlantic Marine Mammal Association 
News Number 9: 9-10. 

Mead, J. G. 1984. Survey of reproductive data for the 
beaked whales (Ziphiidae). Reports of the Interna- 
tional Whaling Commission, Special Issue 6: 91-96. 

Mercer, M. C. 1975. Modified Leslie-DeLury popula- 
tion models of the Long-finned Pilot Whale 
{Globicephala melaena) and annual production of the 
Short-finned Squid {Illex illecebrosus) based upon 
their interaction at Newfoundland. Journal of the 
Fisheries Research Board of Canada 32: 1 145-1 154. 

Moore, J. C. 1966. Diagnosis and distribution of 
beaked whales from the genus Mesoplodon known 
from North American waters. Pages 33-61 in Whales, 
dolphins and porpoises. Edited by K. Norris. 
University of California Press. 

Orr, R. T. 1953. Beaked whale {Mesoplodon) from 
California, with comments on taxonomy. Journal of 
Mammalogy 34: 239-249. 

0ynes, R. P. 1974. Observajoner og merking av brugde 
og hval i Norskehavet i Mail og Juni 1974. 
Fisherinaergens Fors ksfond, Rappoprter nr. 4-1974: 
43-46. Fiskeridirektoratets havforskningsinstitutt, 
Bergen, Norge. 

Price, C. A., and F. M. Farfield. 1985. Abstract. Use of 
satellite data to correlate beaked whale and squid 
distribution with the shelf/ slope thermal front. 
Proceedings, Sixth Biennial Conference on the 
Biology of Marine Mammals, Vancouver, British 
Columbia. 



130 



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Vol. 104 



Reiner, F. 1986. First record of Sowerby's Beaked 
Whale from Azores. Science Report for the Whales 
Research Institute 37: 103-107. 

Saemundsson. B. 1939. Mammals. The zoology of 
Iceland IV, E. E. Munksgaard, Copenhagen 76: 1-52. 

Sergeant, D. E. 1982. Mass strandings of toothed 
whales (Odontoceti) as a population phenomenon. 
Science Report for the Whales Research Institute 
Number 34: 1-47. 

Sergeant, D. £., and H. D. Fisher. 1957. The smaller 
Cetacea of eastern Canadian waters. Journal of the 
Fisheries Research Board of Canada 14(1): 83-115. 

Sergeant, D. E., A. W. Mansfield, and B. 
Beck. 1970. Inshore records of Cetacea for eastern 
Canada, 1949-68. Journal of the Fisheries Research 
Board of Canada 27: 1903 1915. 

Sheldrick, M. C. 1979. Cetacean strandings along the 
coasts of the British Isles 1913-1977. Pages 35-53 in 
Biology of marine mammals: insights through 
strandings. Final Report to the U.S. Marine Mammal 
Commission in Fulfillment of Contract MM7AC020. 



Edited by J. G. Geraci and D. J. St. Aubin. U.S. 

Marine Mammal Commission, Washington, D.C. 

Report Number of MMC-77/ 13; PB 293890. 
True, F. W. 1910. An account of the beaked whales of 

the family Ziphiidae in the collection of the U.S. 

National Museum, 73: 1-89. 
Tyack, P. L. 1987. Abstract. A comparative view of 

mass stranding in cetacea. Page 71 in Seventh Biennial 

Conference on the Biology of Marine Mammals, 

Miami, Florida, December 1987. 
Ulmer, F. A., Jr. 1941. Mesoplodon mirus in New 

Jersey, with additional notes on the New Jersey M. 

densirostris, and a list and key to the ziphioid whales of 

the Atlantic coast of North America. Proceedings of 

the Academy of Natural Science of Philadelphia 93: 

107-122. 
Watson, L. 1981. Sea guide to whales of the world. 

Nelson Canada, Scarborough, Ontario. 302 pages. 



Accepted 10 October 1989 



Status of Stejneger's Beaked Whale, Mesoplodon stejnegeri, 
in Canada* 



J. Houston 



374 Fireside Drive, Woodlawn, Ontario KOA 3M0 

Houston, J. 1990. Status of Stejneger's Beaked Whale, Mesoplodon stejnegeri, in Canada. Canadian Field- 
Naturalist 104(1): 131-134. 

Stejneger's Beaked Whale, Mesoplodon steinegeri, is a medium-sized ziphiid of the North Pacific. The range extends 
from Japan to the North American coast between 50 to 60° N. The species is known only from a few stranded 
specimens, although unsubstantiated sightings have been recorded from Japan and Washington State. The species is 
of no commercial interest and is rare in Canadian waters. 

La baleine a bee de Stejneger, Mesoplodon stejnegeri, est un Ziphiide de taille moyenne du Pacifique nord. Son aire de 
repartition s'etend du Japon a I'Amerique du Nord, entre 50° et 60° de latitude nord. L'espece ne nous est connue que 
par quelques specimens echoues; il semble toutefois qu'elle ait ete apargue au Japon et dans I'Etat de Washington, mais 
ces observations n'ont pas ete confirmees. L'espece ne fait I'objet d'aucune chasse commerciale et est rare dans les eaux 
canadiennes. 

Key Words: Stejneger's Beaked Whale, Mesoplodon stejnegeri, mesoplodon de stejneger, Bering Sea Beaked Whale, 
Cetacea, odontocetes, beaked whales, Ziphiidae, Mesoplodon. 



Stejneger's Beaked Whale, Mesoplodon stej- 
negeri True 1885, was first named on the basis of a 
skull found by Leonhard Stejneger on Bering 
Island, Alaska in 1883. True later included a full 
description in an account of the family (True 1910) 
based on an adult male stranded near Newport, 
Oregon in 1904. Moore (1963, 1966, 1968) 
suggested that the species was the dominant 
subarctic beaked whale of the North Pacific and 
that it can easily be confused with the Splay- 
toothed Beaked Whale {Mesoplodon bowdoini), 
of the South Pacific. 

This species is difficult to identify in the field, but 
any beaked whale in the North Pacific, north of 
50° N latitude, lacking the white beak of Hubbs' 
Beaked Whale {Mesoplodon carlhubbsi) is apt to be 
Stejneger's Beaked Whale (Watson 1981; Mead et 
al. 1982). The only other beaked whale found as far 
north is Cuvier's Beaked Whale {Ziphius cavirostris) 
which has a shorter beak and exposed teeth set on 
the anterior end of the mandible beyond the tip of 
the upper jaw (Leatherwood et al. 1982). 

Stejneger's Beaked Whale has the usual spindle- 
shaped body of the ziphiids (Figure 1) and is of 
medium size. Maximum length is probably 6 m 
with a weight of 1200 kg. The beak is long, the 
fulcate dorsal fin is behind the midline and there is 
no medium notch between the flukes (Leather- 
wood et al. 1982). The flippers are small and there 
are a pair of throat creases typical of beaked 
whales. There is a well-developed keel from the 



dorsal fin to the flukes (Watson 1981). Two 
mandibular teeth which dorsally constrict the 
upper jaw erupt in males. 

As with most other ziphiids, few of these whales 
have been observed alive and the species is known 
mainly from strandings. The colour pattern is not 
well known but appears characteristically to 
include white blotches around the lips, sides of the 
head and neck, and a brown saddle across the 
blowhole (Watson 1981). The animals are grayish 
brown dorsally and lighter on the ventral surface. 
Adults are usually heavily marked with oval white 
scars on the flesh and underbelly and adult males 
demonstrate linear scars which may result from 
intraspecific aggression (Leatherwood et al. 1982; 
Heyning 1984). 

Distribution 

Stejneger's Beaked Whale is known almost 
entirely from records of strandings and from 
stranded specimens, which suggest the species is 
endemic to the subarctic and cold temperate waters 
of the North Pacific (Figure 2) principally between 
50° and 60° N (Moore 1966). 

The species has been recorded from Japan in the 
west and from Alaska to Monterey, California, in 
the east (Moore 1966; Watson 1981; Leatherwood 
et al. 1982). Stranding records exist from 
Vancouver Island (Pike and MacAskie 1969) 
suggesting that the waters off Canada's Pacific 
coast are within the normal range of the species. 



♦Reviewed and accepted by COSEWIC 11 April 1989 — no designation required. 



131 



132 



The Canadian Field-Naturalist 



Vol. 104 




Figure 1 . Stejneger's Beaked Whale, Mesoplodon stejnegeri (redrawn and adapted from 
Watson 1981). 



Protection 

International: All Mesoplodon species are 
included on Appendix II of the Convention on 
International Trade in Endangered Species 
(CITES). Such species are not considered to be 
rare or endangered but could become so if trade 
were not regulated. Listing on Appendix II 
requires management by range states and export 
permits from country of origin for export of parts, 
specimens, or their derivatives. Mitchell (1975) 
Usts the species as one not generally taken or of 
interest to whalers. The species is listed in the Red 
Data Book of the International Union for the 
Conservation of Nature as of "Indeterminate" 
status (Goodwin and Holloway 1972). 

National: Protected by general legislation in 
several countries but no specific provisions are 
known. In Canada, general protection is granted 
under the Fisheries Act and the Cetacean 
Protection Regulations which prohibit whaUng. In 
the United States, general protection is provided 
under the Marine Mammals Protection Act of 
1972 and the Endangered Species Act. 

Population Sizes and Trends 

The species is known only from limited stranding 
records which are few and far between. Only seven 
specimens were recovered between 1910 and 1960 
from Alaska (4), to Vancouver (1), to Washington 
(1) and Oregon (1) [Moore 1966]; one additional 
individual was found in Japan during this same 
period. Since 1960, an additional five strandings 
have been reported (Watson 1981), mostly from the 
Aleutian Islands, but one individual was found near 
Monterey, California (Leatherwood et al. 1982). 
There are unconfirmed reports of sightings in the 
Sea of Japan (Nishimura and Nishiwaki 1964) and 
coastal Washington (Mitchell 1975). 

Although Mitchell (1975) felt that numbers may 
not be as low as sightings and strandings might 



indicate, these whales are undoubtedly rare in 
Canadian waters and have apparently been 
harvested in Japan by Japanese coastal whalers on 
occasion (Mitchell 1975). No other information is 
available on population status. 

Habitat 

The distribution of Stejneger's Beaked Whale 
appears to coincide with the Subarctic current 
systems described by Favorite et al. (1976). The 
distribution may not be as directly related to 
oceanographic parameters as it is to the prey 
species. However, the distribution of its prey 
(squid, pelagic fish) is no doubt related to 
characteristics of the water mass (Favorite et al. 
1976; Watson 1981). 

The species seems to occupy the same subarctic 
niche as Sowerby's Beaked Whale {Mesoplodon 
bidens) in the North Atlantic (Moore 1966; 
Watson 1981) where the latter species also appears 
to be restricted to cooler, offshore waters where 
squid are plentiful (Sergeant and Fisher 1957). 

General Biology 

There is very little information on the natural 
history of the species. Stomach contents of 
specimens taken by Japanese coastal whalers are 
said to consist of squid and pelagic fish (Nishiwaki 
and Kamiya 1959). The species is also reported by 
the whalers to occur in small groups of two to three 
individuals. The largest reported female was 5.3 m 
in length as was the largest recorded male (Mead 
1984). No fetuses or calves have been reported and 
no other morphometric data appears to be 
available. 

Limiting Factors 

Owing to the paucity of information on the 
species, nothing is known of factors which could be 
limiting the population(s). Their solitary habits, 



1990 



HOUSTON: Status of the Stejneger's Beaked Whale 



133 



160 laO 120 '00 80 60 40 20 




Figure 2. Distribution of Stejneger's Beaked Whale. 



distribution and apparent rarity may serve to offer 
some protection from human exploitation. 
Apparently, animals have been taken occasionally 
by Japanese whalers in coastal fisheries (Nishiwaki 
and Kamiyu 1959). The species does not appear to 
have been commercially exploited and the 
occasional removal of a few individuals is unhkely 
to affect the population(s). Commercial exploita- 
tion is not likely, but would probably seriously 
affect existing populations. 

Special Significance of the Species 

The population is generally too small and 
individuals too scarce to be of commercial interest, 
although some species of Mesoplodon may have 
been fished in the 19th Century, or earlier, by 
American and other whalers (Mitchell 1974). The 
blubber and flesh of the species is thought to cause 
diarrhoea (Mitchell 1975; Mead et al. 1982). 

It is improbable that parts or derivatives of the 
species would be involved in international trade. 
The species has not been held in captivity and such 
an event would only come about by chance. 

Evaluation 

Little information is available for assessment of 
population status. Although small groups have 
been observed chasing schools of salmon in the Sea 
of Japan (Nishimura and Nishiwaki 1964). 
Sightings in Canadian waters are few. However, 
the species was known to the Makah Indians of 
Washington who were said to have talked of seeing 
these whales in twos or threes one-half to one mile 
offshore {see Mitchell 1975). Based on the above 
observations, Mitchell (1975) felt the numbers may 
not be small. It is probably rare, at least in 



Canadian waters, but there are no imminent 
threats to its survival here. The species need not be 
considered for COSEWIC status at this time. 

Acknowledgments 

The support of the Deparment of Fisheries and 
Oceans, the Canadian Wildlife Service, and World 
Wildlife Fund in the production of this report is 
gratefully acknowledged. 

Literature Cited 

Favorite, F., A. J. Dodimead, and K. Nasu. 1976. 
Oceanography of the subarctic Pacific Region, 1960- 
71. International North Pacific Fisheries Commission 
Bulletin Number 33. 

Goodwin, H. A., and C. A. Holloway. 1972. Red Data 
Book, Volume I (Mammalia). International Union for 
the Conservation of Nature. Morges, Switzerland. 

Heyning, J. E. 1984. Functional morphology involved 
in intraspecific fighting of the beaked whale, Meso- 
plodon carlhubbsi. Canadian Journal of Zoology 62: 
1645-1654. 

Leatherwood, S., R. R. Reeves, W. F. Perrin, and W. E. 
Evans. 1982. Whales, dolphins, and porpoises of the 
eastern North Pacific and adjacent Arctic waters. U.S. 
Department of Commerce, National Oceanic and 
Atmospheric Administration, National Fisheries 
Service Circular 444. 

Mead, J. G. 1984. Survey of reproductive data for the 
beaked whales (Ziphiidae). Reports of the Interna- 
tional Whaling Commission, Special Issue 6: 91 96. 

Mead, J. G., W. A., Walker, and W. J. Houck. 1982. 
Biological observations on Mesoplodon carlhubbsi 
(Cetacea: Ziphiidae). Smithsonian Contributions to 
Zoology Number 344. 

Mitchell, E. 1974. Conservation of smaller whales, 
dolphins and porpoises. Pages 72 79 in Wildlife 
Today. Edited by N. Sitwell. Tom Stacey Limited, 
London, England. 



134 



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Vol. 104 



Mitchell, E. 1975. Porpoise, dolphin and small whale 
fisheries of the world. International Union for the 
Conservation of Nature, Mongraph Number 3. 

Moore, J. C. 1963. Recognizing certain species of beaked 
whales of the Pacific Ocean. American Midland 
Naturalist 70: 396-428. 

Moore, J. C. 1966. Diagnoses and distributions of 
beaked whales of the genus Mesoplodon known from 
North American waters. Pages 32-61 in Whales, 
dolphins and porpoises. Edited by K. S. Norris. 
University of California Press, Berkeley, California. 

Moore, J. C. 1968. Relationships among the living 
genera of beaked whales with classifications, diagnoses 
and keys. Fieldiana 53: (4): 209-298. 

Nishimura, S., and M. Nishiwaki. 1964. Records of the 
beaked whale Mesoplodon stejnegeri from the Japan 
Sea. Bulletin of the Seto Marine Biology Laboratory 12: 
323-334. 

Nishiwaki, M., and T. Kamiya. 1959. Mesoplodon 
stejnegeri from the coast of Japan. Science Reports of 
the Whales Research Institute 14: 35-48. 



Pike, G. C, and I. B. MacAskie. 1969. Marine 
mammals of British Columbia. Fisheries Research 
Board of Canada Bulletin 171. 

Sergeant, D. E., and H. D. Fisher. 1957. The smaller 
cetacea of eastern Canadian waters. Journal of the 
Fisheries Research Board of Canada 14: 83-115. 

True, F. W. 1885. Contributions to the history of the 
Commander Islands. Number 5. Description of a new 
species of Mesoplodon, M. stejnegeri, obtained by Dr. 
Leonard Stejneger in Bering Island. Proceedings of the 
U.S. National Museum 8: 584-585. 

True, F. W. 1910. An account of the beaked whales of 
the family Ziphiidae in the collection of the United 
States National Museum. Bulletin of the U.S. National 
Museum 73: 1-89. 

Watson, L. 1981. Sea guide to whales of the world. 
Nelson Canada Limited, Scarborough, Ontario. 

Accepted 10 October 1989 



Status of True's Beaked Whale, Mesoplodon mirus, in Canada* 

J. Houston 

374 Fireside Drive, Woodland, Ontario KOA 3M0 

Houston, J. 1990. Status of True's Beaked Whale, Mesoplodon mirus, in Canada. Canadian Field-Naturalist 104(1): 
135-137. 

True's Beaked Whale, Mesoplodon mirus, is a medium-sized cetacean of the family Ziphiidae found in the temperate 
North Atlantic and the southwestern Indian Ocean. Known mainly from stranded specimens, the species is probably 
pelagic, but occasionally seen in coastal waters. It is not known to have been commercially exploited or taken in other 
fisheries operations. The species is probably rare in Canadian waters. 

La baleine a bee de True, Mesoplodon mirus, est un cetace de taille moyenne appartenant a la famille des Ziphiides et 
frequentant les eaux temperes de I'Atlantique nord et du sud-ouest de I'Ocean Indien. Les donnees disponibles sur 
I'espece proviennent d'individus echoues; probablement pelagique et rare, elle evite les zones cotieres. On ne sait pas si 
elle a fait I'objet d'une exploitation commerciale ou si elle a ete capturee au cours d'autres operations de peche. L'espece 
est probablement rare dans les eaux canadiennes. 

Key Words: True's Beaked Whale, Wonderful Beaked Whale, Mesoplodon mirus, baleine a bee de True, Cetacea, 
odontocetes, ziphiids, beaked whales, Mesoplodon, North Atlantic. 



True's Beaked Whale, Mesoplodon mirus True 
1913, is a medium-sized ziphiid, the maximum 
length observed being between 5 and 5.5 m with an 
average weight of 1300 kg (Watson 1981; 
Leatherwood and Reeves 1983). The body is 
distinctive (Figure 1 ) and similar to that of Cuvier's 
Beaked Whale {Ziphius cavirostris). The head is 
small with a bulging forehead and short neck. The 
two teeth which erupt in adult males are found near 
the tip of the mandible. The body is chunky in the 
midriff, tapering towards the tail. There is a slight 
depression behind the blowhole. The flippers are 
small and positioned low and forward similar to 
those of Cuvier's Beaked Whale. Behind the small, 
curved dorsal fin, is a pronounced dorsal ridge. 
The flukes have a small median notch unlike those 
of other beaked whales {see Watson 1981; 
Leatherwood and Reeves 1983). 

The species is usually bluish-gray to dark gray 
on the back, lighter gray on the sides and gray 
ventrally. Lighter spots or blotches are often found 
on the throat and anal regions (Watson 1981; 
Leatherwood and Reeves 1983). Scarring, some of 
which is probably from the teeth of other males, is 
common in adult males (Heyning 1984). 

As with other ziphiids, the species is rarely 
observed and is known mostly from strandings. 
Apparently rare, at least not common in Canadian 
waters, the status of the species in Canada is of 
interest to the Committee on the Status of 
Endangered Wildlife in Canada (COSEWIC). This 
report was prepared for the Committee's 
consideration. 



Distribution 

The species is known only from a few strandings 
and, until recently, was thought to be restricted to 
the temperate North Atlantic (Figure 2). In 1959, a 
specimen was found on the South African coast 
(Talbot 1960) and several others have been recorded 
since (McCann and Talbot 1963; Ross 1969). The 
occurrence of M. mirus in the South Atlantic may 
represent a geographically separate stock, or that 
the range may be wider than previously thought 
(Leatherwood and Reeves 1983). 

In the North Atlantic, strandings have been 
recorded from Florida to Nova Scotia in the west, 
and on the west coast of the British Isles in the east 
(Sergeant and Fisher 1957; Moore 1966; Brown 
1975). Most strandings have been recorded along 
the North American coast, the centre of 
distribution is about 38°N (Moore 1966). Moore 
(1966) considered Mesoplodon mirus to be 
allopatric with Sowerby's Beaked Whale 
{Mesoplodon bidens), which occurs more 
frequently in European waters and further to the 
north. 

Protection 

International: The species, along with other 
members of the genus is listed on Appendix II of 
the Convention on International Trade in 
Endangered Species (CITES). No other interna- 
tional agreements refer to the species. Mitchell 
(1975a) considered the species to be of Indetermi- 
nate Status and it is listed as such in the Red Data 
Book (Goodwin and Holloway 1972). 



♦Reviewed and Accepted by COSEWIC 11 April 1989 — no designation required. 

135 



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The Canadian Field-Naturalist 



Vol. 104 




Figure 1 . True's Beaked Whale, Mesoplodon mirus (redrawn and adapted from Watson 
1981). 



National: The species is protected by general 
legislation in several countries but no specific 
provisions are known. In Canada, general 
protection is provided under the Cetacean 
Protection Regulations promulgated under the 
Fisheries Act. In the United States, general 
protection is provided under the Marine Mammals 
Protection Act and the Endangered Species Act. 

Population Sizes and Trends 

As with other ziphiids that are known 
principally from stranding records, it is not 
possible to determine population numbers. They 
have not been subject to commercial exploitation 
and it is unknown if the numbers have changed 
over the last century (Mitchell 1975a, 1975b). It 
may be more widespread than previously thought 
as the South African strandings are probably 
indicative of a separate stock (McCann and Talbot 
1963; Ross 1969; Watson 1981). 

Only one stranding has been reported in 
Canada, on Cape Breton Island in 1938 (Moore 
1966), and this is the most northerly record from 
the western North Atlantic. Other North American 
strandings are few (eight) and have occurred along 



the United States coast from Maine to Florida. In 
the eastern North Atlantic four records exist, one 
from the Orkney Islands and three from the west 
coast of Ireland (Moore 1966; Brown 1975; 
Watson 1981). Since this is a pelagic species, it is 
rarely encountered alive and no verified sightings 
at sea are known. 

Habitat 

As the species is rarely observed, there is no 
information on its habitat, except for the limited 
distributional data provided by strandings. The 
species' diet appears to consist of squid and deep- 
water fish (Mitchell 1975b; Watson 1981), thus 
True's Beaked Whale probably inhabits the open 
ocean rather than coastal areas, as do most other 
ziphiids. 

General Biology 

Very little is known about the life history of the 
species. Most information comes from stranded 
individuals. They appear to eat squid and pelagic 
fish (Mitchell 1975b; Watson 1981) and are Hkely 
solitary. Calves are probably born in the spring and 
are about 2 m long at birth (Brimley 1943; Moore 
1966; Watson 1981). 



160 140 120 '00 80 60 40 20 




FiCitiRi- 2. Distribution olTruc's Beaked Whale, Mesoploilon mirus. 



1990 



HOUSTON: Status of True's Beaked Whale 



137 



The largest female reported was 5.1 m and the 
largest male 5.3 m; the shortest calf measured was 
2.3 m and the largest fetus 1.1m (Mead 1984). 
Aside from some meager data on ovarian and 
testicular weights {see Mead 1984), no other 
morphometric information appears to be 
available. 

Limiting Factors 

None known. The species is not known to have 
been exploited commercially or taken incidentally 
in other fishing operations (Mitchell 1975a, 
1975b). Its rarity and solitary lifestyle probably 
help to maintain natural protection. As the food 
species appear to be pelagic, these whales probably 
avoid coastal areas which are more apt to be 
subject to pollution. The species appears to be shy 
of ships and dives on approach (Watson 1981). 

Special Significance of the Species 

True's Beaked Whale is not known to be, or to 
have been, subject to exploitation; no specimens 
have been recorded in captivity. 

This beaked whale is easily confused with 
Cuvier's Beaked Whale (Leatherwood and Reeves 
1983). Identification of all beaked whales is 
difficult and often requires the assistance of an 
expert. 

Evaluation 

The species may be more widely distributed than 
previously thought but is probably nowhere locally 
abundant in coastal areas. No confirmed sightings 
have been recorded (Watson 1981) and strandings 
on the Atlantic coast of Canada have been few and 
far between. The species is probably naturally rare 
in Canadian waters, but is not under any imminent 
threat; no COSEWIC status designation is 
required at this time. 

Acknowledgments 

The support of the Department of Fisheries and 
Oceans, the Canadian Wildlife Service, and the 
World Wildlife Fund (Canada) are gratefully 
acknowledged. 



Literature Cited 

Brimley,H. H. 1943. A second specimen of True's beaked 
whale, Mesoplodon mirus True, from North Carolina. 
Journal of Mammalogy 24: 199-203. 

Brown, S.G. 1975. Relation between stranding mortality 
and population abundance of smaller Cetacea in the 
northeast Atlantic Ocean. Journal of the Fisheries 
Research Board of Canada 32(7): 1095-1099. 

Goodwin, H. A., and C. W. HoUoway. 1972. Red Data 
Book, Volume I (Mammalia). International Union for 
the Conservation of Nature and Natural Resources, 
Morges, Switzerland. 

Heyning, J. E. 1984. Functional morphology involved in 
intraspecific fighting of the beaked whale, Mesoplodon 
carlhubbsi. Canadian Journal of Zoology 62: 
1645-1654. 

Leatherwood, S., and R. R. Reeves. 1983. The Sierra 
Club handbook of whales and dolphins. Sierra Club, 
San Francisco, California. 

McCann, C, and F. H. Talbot. 1963. The occurrence of 
True's Beaked Whale in South African waters, with a 
key to South African species of the genus. Proceedings 
of the Linnean Society, London 175: 137-144. 

Mead, J. G. 1984. Survey of reproductive data for 
beaked whales (Ziphiidae). Reports of the International 
WhaUng Commission, Special Issue 6: 91-96. 

Mitchell, E. 1975a. Porpoise, dolphin and small whale 
fisheries of the world. International Union for the 
Conservation of Natural Resources, Morges, Switzer- 
land, Monograph Number 3. 

Mitchell, E. D. ^j/^or. 1975b. Report of the Meeting on 
Smaller Cetaceans, Montreal, April 1-11, 1974, 
Subcommittee on Small Cetaceans Scientific Commit- 
tee, International Whaling Commission. Journal of the 
Fisheries Research Board of Canada 32(7): 889-983. 

Moore, J. C. 1966. Diagnoses and distributions of 
beaked whales of the genus Mesoplodon known from 
North American waters. Pages 32-61 in Whales, 
dolphins and porpoises. Edited by K. S. Norris. 
University of California Press, Los Angeles, California. 

Ross, D. J. B. 1969. Evidence for a southern breeding 
population of True's beaked whale. Nature 222: 585. 

Sergeant, D. E., and H. D. Fisher. 1957. The smaller 
cetacea of eastern Canadian waters. Journal of the 
Fisheries Research Board of Canada 14: 83-1 15. 

Talbot, F. H. 1960. True's beaked whale from the 
southwest coast of South Africa. Nature 186: 406. 

Watson, L. 1981. Sea guide to whales of the world. 
Nelson Canada Limited, Scarborough, Ontario. 

Accepted 10 October 1989 



Status of the Ringed Seal, Phoca hispida, in Canada* 

Michael C. S. Kingsley 

Department of Fisheries and Oceans, Institut Maurice-Lamontagne, 850 Route de la Mer, Mont-Joli, Quebec 
G5H 3Z4 

Kingsley, Michael C. S. 1990. Status of the Ringed Seal, Phoca hispida, in Canada. Canadian Field-Naturalist 
104(1): 138-145. 

The Ringed Seal, Phoca hispida, is a small seal ubiquitous in ice-covered seas in the northern hemisphere. Although it 
has been hunted for its meat, blubber, and hide by the native people of the north for millennia, and is also the mainstay 
of the Polar Bear, its populations remain stable, varying only in response to variations in the weather. It is adaptable to 
human disturbance of many kinds. 

Le phoque annele, Phoca hispida, est un petit phoque que Ton trouve dans toutes les mers couvertes de glace de 
I'hemisphere nord. Bien que Tespece ait ete chassee pour sa chair, sa graisse et sa peau par les autochtones du Nord 
pendant des millenaires, et qu'elle constitue egalement la principle proie de Fours blanc, sa population demeure stable, 
ne variant qu'en fonction des conditions atmospheriques. Ce phoque s'adapte tres bien aux perturbations 
anthropiques de toutes sortes. 

Key Words: Ringed Seal, Phoca hispida, Phocidae, seals, phoque annele, status, Canada. 



Phoca hispida Schreber 1 775, the Ringed Seal of 
the Arctic, is the smallest seal, less than half the 
weight of other phocids (Stirling 1977). It is dark 
gray, with lighter-gray rings on the back and flanks 
and a paler gray ventral surface (Figure 1). Mature 
males develop a black facial mask in the spring. 
Adult females in the western Canadian Arctic 
average 122 cm long, males 136 cm (Smith 1987). 
Alaskan seals are smaller (Frost and Lowry 1981). 

The single pup is born about 69 cm long, 
weighing about 3.5 kg (Smith 1987: 37), and clad in 
a long, white lanugo. The first-year coat which 
replaces the lanugo 20 to 30 days after birth is 
rather longer and thicker than the typical adult 
coat, and less strongly marked. This coat was 
esteemed in the fur trade as the "silver jar" 
(McLaren 1958a) and was for decades a major 
economic resource for Arctic maritime people 
(Smith and Taylor 1977). 

Distribution 

The Ringed Seal is ubiquitous in ice-covered 
seas of the northern hemisphere (King 1983); of the 
Arctic seals, it is uniquely adapted to living and 
breeding in fast ice, in which it maintains breathing 
holes throughout the winter (Stirling 1974, 1977). 

Apart from certain physically separated groups, 
the Ringed Seals of the circumpolar Arctic form a 
continuum (Phoca hispida hispida). Marine 
populations are found in the Sea of Okhotsk 
(Phoca hispida ochotensis) and the northern parts 
of the Baltic Sea (Phoca hispida hotanica) and 



land-locked sub-species are found in Lake Saimaa 
in Finland (Phoca hispida saimensis), and Lake 
Ladoga (Phoca hispida ladogensis) in the USSR 
(Frost and Lowry 1981). Closely related species are 
found in the Caspian Sea (Phoca caspica) and 
Lake Baikal (Phoca sibirica) (Frost and Lowry 
1981). 

The Canadian range for the Ringed Seal is 
depicted in Figure 1. Studies on population 
ecology associated with intense native harvests 
were carried out in south-west Baffin island and 
Foxe Basin in the 1950s (McLaren 1958a), in 
south-east Baffin Island in the late 1960s (Smith 
1973b), and in the western Arctic in the 1970s 
(Smith 1987). Site-specific studies, components of 
assessments of the expected environmental 
impacts of proposed mines, shipping, or pipelines, 
have been made in the Northwest Passage (Finley 
1976; Finley and Johnson 1977; Smith et al. 1978; 
Smith and Hammill 1980a; Kingsley et al. 1985) 
and in the Beaufort Sea, Amundsen Gulf and 
Coronation Gulf (StirUng et al. 1977, 1982; 
Alliston and McLaren 1981; Kingsley 1984, 1986). 
Less is known about density and distribution in the 
permanent polar pack of the Arctic basin. 

Protection 

International Protection Measures: There are 
no international agreements or conventions 
intended to protect Ringed Seals. The Interna- 
tional Agreement on the Conservation of Polar 
Bears and their Habitat protects the ecosystems of 



♦Reviewed and accepted by COSEWIC I I April 1989 no designation required. 



138 



1990 



KiNGSLEY: STATUS OF THE RINGED SEAL 



139 




Figure 1. The Ringed Seal, Phoca hispida (drawn by J. Venables; courtesy Alaska 
Department of Fish and Game). 




Figure 2. The range of the Ringed Seal in Canada. 



which Polar Bears {Ursus maritimus) are a part 
with special attention, inter alia, to feeding sites 
(Stirling 1985). This implies a measure of 
protection for Ringed Seals and their habitat. 

National Protection Measures: Seals in the 
Canadian Arctic are protected under the Seal 
Protection Regulations made under the Fisheries 
Act. These permit any resident to take seals for 
food for himself, his family or his dogs, or to sell or 



trade seal meat to a resident or a traveller for the 
same purpose (Canada Department of Fisheries 
and Oceans CRC 1978). There are no restrictions 
on the sale or barter of seal skins produced by this 
harvest. There is a provision for sport sealing 
hcences, which allow the holder to take two seals. 
Less than 100 such hcences are sold annually. 

Vulnerable species are gregarious, migratory or 
valuable. The Ringed Seal in the Canadian Arctic 



140 



The Canadian Field-Naturalist 



Vol. 104 



is protected de facto by its wide, uniform 
distribution, its sedentary habits, and its low value. 
Seal skins were worth $25 to $30 in the mid 1970s 
(Smith 1979), but owing to the protest against the 
northwest Atlantic Harp Seal {Pagophilus 
groenlandicus) harvest, they are now of little value 
in the fur trade (Hertz and Kapel 1986). 

In Alaska, all marine mammals come under the 
United States Marine Mammal Protection Act of 
1972, which restricts harvest to coastal natives for 
subsistence or the production of authentic 
handicrafts, and restrains regulation of native 
harvest of marine mammals unless a population is 
found to be depleted. 

Population Densities, Sizes and Trends 

The Ringed Seals in the circumpolar Arctic form 
a genetic continuum; separate populations have 
not been defined for any areas of the Canadian 
Arctic, nor have principles been established on 
which such a definition might be attempted. Little 
is known about individual movements but returns 
from tagged seals have shown that sub-adults may 
travel long distances (Smith 1967). 

Numbers can be estimated by shipborne line 
transect survey, and corrected for submerged 
animals (McLaren 1961; Eberhardt 1978; Eber- 
hardt et al. 1979). Density indices may be based on 
satellite or high altitude aerial counts of the 
drained areas around seal holes (Finley 1979; 
Digby 1984) or on the frequency of vocalizations 



(StirUng 1973; Stirhng et al. 1983; Calvert and 
Stirling 1985). Searching the ice in winter or spring 
with dogs which can smell out lairs and breathing 
holes, has been used (Smith and Stirling 1978; 
Smith and Hammill 1980b), but is slow and 
expensive. Under-snow lairs may be detected by 
airborne infra-red sensing (MGSK, unpublished). 
Aerial surveys flown at the summer peak of open- 
air haul-out can give rapid and cheap information 
on the relative densities of seals over wide areas. 
This has been the commonest recent method and 
many hours of such surveys have been flown 
(Table 1). There are problems in interpreting such 
density estimates, because at that time of year, 
seals are probably starting to move from their 
wintering grounds. In particular, sub-adults which 
wintered outside the fast-ice breeding areas invade 
the fast ice to find haul-out sites, and high local 
densities can occur behind ice edges. The fraction 
of seals hauled out is not accurately known. A 
correction factor of 2 was applied to survey data by 
Smith (1973a, 1975); peak haul-out has been 
estimated at 70 to 80% (Finley 1979; Smith and 
Hammill 1981). In general, high latitudes and 
multi-year ice have lower densities (Smith et al. 
1979; Kingsley et al. 1985). 

Seal densities have been measured by studies 
local in scope and short in duration. There is little 
information on trends. Repeated surveys in the 
Beaufort Sea from 1974 through 1979 gave highly 
variable estimates of total numbers (Stirling et al. 



Table 1. Estimates of numbers and densities of Ringed Seals in the Canadian Arctic 



Area 



Period 



Population Density 
Estimate 
(visible seals) Range (/km^) 



Reference 



Beaufort Sea 
Amundsen Gulf 
Northern Amundsen Gulf 
Prince Albert Sound 
Minto Inlet 
Prince Albert Sound 
Prince Albert Sound 
(jsntrai High Arctic 

(Polar Gas Region 1) 
High Arctic (173°30'-78° 10' 
Hudson Bay 
James Bay 
Baffin Bay pack ice 
East Baffin Island — fast ice 
Eastern Arctic (north, east and 

South Baffin Island, 

Foxe Basin, Hudson Bay 

and Strait, and 

Ungava Bay) 
Home Bay (East Baffin Island) 
Hoare Bay (southeast Baffin 

Island) 
Cumberland Sound 



1974-1979 


40 000 


0.1 ■ 


■0.7 


Stirling et al. 1982 


1972 




0.9 ■ 


- 1.1 


Smith 1973a 


1981-1985 


52 000 


1.6 ■ 


-3.1 


Kingsley 1986 and unpublished 


1972 




2.12 




Smith 1973a 


1972 




0.18 




Smith 1973aa 


1981-1985 


18 000 


2.0 ■ 


-3.5 


Kingsley 1986 and unpublished 


1988 


49 000 


2.0 ■ 


-3.5 


Kingsley and Byers, in press 


1975 




0.58- 


■ 1.54 


Finley 1976 


1980-1981 


90 000 


0.06- 


- 1.2 


Kingsley et al. 1985 


1974 


203 000 






Smith 1975; Davis et al. 1980 


1974 


28 000 






Smith 1975; Davis et al. 1980 


1978-1979 


420 000 


1.39 




Finley etal. 1983 


1978-1979 


67 000 
947 000 


0.86- 


-2.3 


Finley etal. 1983 
McLaren 1958b 



1969 
1969 

1969 



33 000 
1 7 000 

27 500 



Smith 1973b 
Smith 1973b 

Smith 1973b 



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KiNGSLEY: STATUS OF THE RINGED SEAL 



141 



1982), but the interpretation of these results is 
complicated by year-to-year variation in the ice 
cover when the surveys were flown. Surveys off the 
Tuktoyaktuk Peninsula from 1981 through 1984 
gave rather constant estimates of total numbers 
(Kingsley 1986), and repeated surveys in Amundsen 
Gulf (Kingsley 1986) and a once-repeated extended 
survey in the high Arctic (Kingsley et al. 1985) also 
gave fairly stable total estimates. 

Fluctuation in the price paid for seal skins, the 
number of dog teams, the availabihty of wage 
employment, and the abundance of other meat such 
as caribou have caused the intensity of seal hunting 
to change over the last 20 years, and resource 
extraction activity has increased at the same time. 
There is yet no indication that Ringed Seal densities 
have altered in response to these changes. The 
numbers of Ringed Seals appear to be stable, with 
some fluctuation in response to episodes of 
unfavourable weather. 

Habitat 

The preferred habitat of the Ringed Seal is in 
areas which freeze every winter to stable fast ice, but 
which have a reliable open-water season. In 
extended aerial surveys of the Canadian high Arctic, 
densities were found to be much lower in areas in 
which there was little summer break-up (Smith et al. 
1979; Kingsley et al. 1985). Information on density 
and distribution in the Arctic basin is not available. 
Water 50 to 150 m deep appears to be preferred. 
Surveys in the Canadian Beaufort Sea showed 
Ringed Seals to be less abundant in shallow water 
off the Tuktoyaktuk Peninsula (Stirling et al. 1982), 
and, in the high Arctic, less abundant in water 
deeper than 175 m. In the coastal waters of 
southwest Baffm Island, Ringed Seals were more 
numerous along complex highly-indented coasthnes 
than simple ones (McLaren 1958a). 

The preferred breeding habitat is reliable fast ice 
which will stay in place until pups are weaned. 
Breeding adults occupy this habitat and apparently 
maintain territories in it, aggressively excluding sub- 
adults (Stirling 1973; Smith and Hammill 1981; 
Smith 1987). Ringed Seals in Baffin Bay are inferred 
(Finley et al. 1983) to breed in the pack ice. Sub- 
adults winter in shear zones and areas of 
unconsolidated ice not suitable for breeding, 
sometimes in high densities and poor condition (M. 
Hammill, Department of Fisheries and Oceans, 
Lamontagne Institute, Mont-Joli, Quebec; unpub- 
lished data), and they are subject to intense 
predation by bears (Stirling and Archibald 1977). 

General Biology 

The female Ringed Seal matures reproductively 
at about 6 yrs, and age-specific pregnancy rates 
reach 0.88 to 0.90 by the 10+ yr age class (Smith 



1987). Maturity may occur as early as 4+ yrs, but 6+ 
looks from the demographic tables hke a normal 
age. A female may bear one pup a year. There is no 
data on intra-uterine mortahty. In late winter, 
females dig enlarged lairs under snowdrifts in which 
they bear their pups in early to mid-April (Smith 
and Stirling 1975). One can infer from descriptions 
of behaviour (e.g. Smith and Stirling 1975) that they 
feed during the 30-day lactation. 

Episodes of reproductive failure, lasting more 
than one year, have occurred in the western 
Canadian Arctic. Ovulation rates dropped 
temporarily to less than half normal values in the 
western Arctic in 1973 to 1975 (Stirling et al. 1977; 
Smith and Stirling 1978; Smith 1987), presumably 
owing to a winter and following summer of heavy ice 
cover. At Sachs Harbour in 1987, ovulation rates 
were low and the 1984 to 1987 cohorts were 
infrequent in collections in 1987 and 1988 (Kingsley 
and Byers 1988, 1989). 

The males are larger than the females and, in the 
breeding season, give off a strong acrid odour 
associated with the apocrine glands in the face 
(Smith 1987). Ringed Seals are thought to be 
polygynous; males probably estabUsh and hold 
underwater territories that include the birth- and 
haul-out lairs of several females. Females ovulate 
and mate at about the time of weaning in mid- to late 
May, but the blastocyst does not implant until 
August or September; active pregnancy lasts on 
average for about 240 days (Smith 1987). 

Movements: The Ringed Seal in Canada is 
pelagic in open water in summer, not hauling out on 
land except under imminent threat from aquatic 
predators (T. Smith, Department of Fisheries and 
Oceans, Arctic Biological Station, Ste-Anne-de- 
Bellevue, Quebec; personal communication). As ice 
forms, consolidates, and thickens, seals move into it 
to establish home ranges (Burns et al. 1980), 
maintaining breathing holes by scratching away the 
ice with their foreclaws. Splashes and exhalations 
form an icy dome over the top (Smith and StirUng 
1975). In mild weather in early winter, seals may 
enlarge holes and haul out in the open. As snow 
cover develops, seals maintain holes, or make fresh 
ones, under snowdrifts, and dig haul-out lairs within 
the drifts. Temperatures within these lairs are 
between -2° and -8°C (M. Kingsley, unpublished 
data) at ambient temperatures of -35° C. 

From mid-May on, seals haul out in the open air 
at holes or on the edges of narrow cracks to bask in 
the sun and moult; they feed little at this time, and 
lose weight (McLaren 1958a; Smith 1987). Haul- 
outs may last up to 40 h, but are usually shorter 
(Smith and Hammill 1980b). The longest haul-out 
recorded for an individual was 40.5 h [Smith and 
Hammill 1981: Table 5 (note that this was the tail 
of a skewed distribution)]. More seals haul out in 



142 



The Canadian Field-Naturalist 



Vol. 104 



the middle of the day (Finley 1979; Smith and 
Hammill 1981) with a daily peak of haul-out at 
1500 h at the start of the haul-out season, becoming 
less clearly defined later. As the ice breaks up, seals 
move to haul out at holes and cracks behind 
retreating ice edges, and very high local 
aggregations result: 15 seals may be seen ringing a 
single hole, or 200 along a kilometre of crack. 
Ringed Seals will only haul out at holes or narrow 
cracks in fast ice or in the middle of very large floes, 
not on the edge of wide leads or ice floes (M. 
C. S. K., unpublished). As the ice rots and breaks 
up, they become aquatic and start to feed 
intensively to regain condition lost in the moulting 
fast. Ringed Seals may aggregate in summer into 
large groups, apparently feeding on local 
concentrations of food (Smith 1987). 

Maturing subadults move into the breeding 
areas and establish home ranges, probably staying 
in the same area throughout their reproductive 
lives. Longevity of 43 yrs has been observed, but 
few seals over 20 yrs occur in the wild (Smith 1987). 

Behaviour and Adaptability: The Ringed Seal is 
asocial. Apart from loose feeding aggregations in 
summer open water, adult Ringed Seals are 
solitary, and live in a state of mild mutual 
repulsion. Groups of seals hauled out at holes are 
rarely large, even in areas of high density (Finley 
1976; Kingsley 1989), and aggression between seals 
at haul-out sites is frequent (Smith and Hammill 
1985). 

Vocalization in Ringed Seals is not well 
developed or frequent. Wide-band underwater 
barks and yelps may be heard about breeding time, 
associated probably with a short-range antago- 
nism, but they are infrequent and quiet. At other 
times of the year, Ringed Seals seem to be less 
vocal (Stirlng 1973; Stirling et al. 1983). 

Ringed Seals, especially experienced adults, are 
alarmed by disturbances that are new, close or 
loud. They leave lairs they are hauled out in at the 
sound of footsteps on the ice or at the sound of a 
landing helicopter (Burns and Kelly 1982). They 
dive when aircraft, especially helicopters, fly low 
over them. Densities were lower close to Eskimo 
villages in Alaska than further away, probably 
because of hunting and snowmobile traffic (Burns 
et al. 1980), and seals were displaced from the 
immediate area of seismic activity (Burns and 
Kelly 1982). However, they easily get used to a 
wide spectrum of human activities and disturban- 
ces which are repeated and not intrusive. 
Investigations in the Canadian and Alaskan 
Beaufort Sea failed to show any lower seal 
densities in areas affected by through-the-ice 
seismography or other industrial activity (Alliston 
1980; Burns and Kelly 1982; Kingsley 1984; Frost 
et al. 1985; Kingsley 1986) and the abundance and 



availability of seals in winter leads, due to drilling 
rigs in the Canadian Beaufort Sea, attract Polar 
Bears (Stirling 1988). Hauled-out seals will become 
accustomed to repeated passage of snowmobiles 
and the density of Ringed Seals off the end of the 
runway at Resolute, Northwest Territories, was 
not lower than elsewhere in spite of heavy aircraft 
traffic and frequent snowmobile passages (Calvert 
and Stirling 1985). 

Food Habits and Feeding: The Ringed Seal is an 
opportunistic feeder on pelagic organisms. Sessile 
benthos is not prominent in the diet, but demersal 
invertebrates are taken (Frost and Lowry 1984). In 
the ice season, the Arctic Cod {Boreogadus saida) 
is the mainstay in most areas, but other fishes such 
as Saffron Cod {Eleginus gracilis) are important at 
some times or in some areas (Lowry et al. 1980). In 
the open-water season, invertebrates are equally 
important. Invertebrate diet composition varies 
from area to area; mysids {Mysis oculata), and 
amphipods, especially Parathemisto libellula, are 
generally important, but may be displaced by other 
locally more abundant species (Lowry et al. 1980; 
Smith 1987). 

Limiting Factors 

This seal is limited by its dependence on ice. In 
southerly waters it is exposed to competition from 
other seals, predation by Killer Whales (Orcinus 
orca) in open water or Walrus in unconsolidated 
ice, and breeding failure due to unstable ice or to 
snow cover inadequate to protect the newborn pup 
from the weather or predators. The species has 
been generally studied in areas of abundance, and 
its interactions with other species at the edges of its 
range are not well known. 

Limits on Density: Territoriality among the 
adult seals limits the density in the fast ice breeding 
habitat (Stirling et al. 1981). It is not known what 
determines the size of the home range that a 
breeding female will try to maintain, nor whether 
seals behave in the same way when breeding on 
pack-ice; other pack-ice seals are seldom territorial 
(Stirling 1983). The ultimate limiting factor is 
thought to be food (McLaren 1958a). Within the 
Arctic ice range of the Ringed Seal, density and 
population structure respond to local variations in 
ice chronology (Stirling et al. 1981). Polynya 
localities where Walrus {Odobenus rosmarus) live 
all year are unsuitable for Ringed Seal because 
Walrus probably prey on any Ringed Seals they 
can catch and drive them out of the area (Fay I960; 
Lowry and Fay 1984). 

Mortality Factors: Four causes of death 
dominate the population dynamics of the Ringed 
Seal. These are hunting by humans, hunting by 
Polar Bears, predation on pups by Arctic Foxes 



1990 



KiNGSLEY: STATUS OF THE RINGED SEAL 



143 



(Alopex lagopus), and intra-specific intolerance 
(Smith 1987). Predation by gulls and Ravens 
{Corvus corax) on newborn pups exposed by lack of 
a covering snowdrift has been recorded (Kumlien 
1879: cited in McLaren 1958a; Lydersen and Smith 
1989) and may impose a limit on distribution, but is 
negligible in most of the species's range. Walrus are 
said to prey on Ringed Seals, which shun areas 
where they regularly occur; this probably has only a 
minor population effect, but may limit distribution 
(Fay 1960; Lowry and Fay 1984). 

The Ringed Seal was the mainstay of the recent 
marine Eskimo culture. It fed man and dog, and lit 
and heated the snowhouse and cooked the food. Its 
hide covered kayaks, and made spring boots, 
mittens, and parkas. More recently, seal pelts, 
especially the "silver jar" from the newly-moulted 
pup, were of value in the commercial fur trade. The 
price of seal skins, the number of dog teams, the 
availability of wage employment, and the 
abundance of other meat such as caribou have all 
varied in recent decades, and affected the intensity of 
seal hunting. The annual kill has never been exactly 
known; a partial record showed it to be of the order 
105 seals/yr (Smith and Taylor 1977; Smith 1987); 
this is probably an underestimate even of the fur 
harvest and would not include those seals used only 
for food. Increasing preservationist influence has 
destroyed the market in sealskins (Smith 1987), and 
the native harvest is now much reduced (Stewart et 
al. 1986). Young seals of the year are taken for food, 
and their skins are still used for making spring 
mittens and boots and for handicrafts. A larger 
harvest is taken for dog food; Polar Bear sport hunt 
regulations require that dog teams be used, so the 
number of dogs has increased in recent years, and 
they are fed largely on seal. The usual method of 
hunting seals is to shoot them when they are hauled 
out on the ice in the early summer or from small 
boats in the open water. Because skins are no longer 
traded and because seal hunting is often regarded as 
a casual pastime, the total harvest of seals is difficult 
to estimate accurately, and there are no reliable 
statistics. Heavy hunting close to communities may 
locally reduce density (Burns et al. 1982), but density 
varies in short range even in unhunted areas. 

The Polar Bear is an obligate carnivore. Its diet is 
composed of marine mammals, mostly Ringed 
Seals. It hunts in winter most successfully in ice-edge 
and shear-zone areas which tend to be densely 
inhabited by naive sub-adult seals, but is less 
successful in catching breeding adults in the fast ice. 
Bears were seen to catch one seal about every five 
days (Stirling 1974). A 200 kg bear (Kingsley 1979) 
has an estimated BMR of 3.7 Mcal/d; a 25 kg seal 
with a caloric density of 5 Meal/ kg (Stirling and 
McEwan 1975) yields 125 Meal, i.e., about 34 days' 
maintenance. Allowing for wastage, assimilation, 



and the difference between average and basal 
metabolism by a factor of 4 gives an annual average 
requirement of 1 seal/ 8.5 days. If there are about 
15 000 to 20 000 Polar Bears in the Canadian Arctic 
(I. Stirling, Canadian Wildlife Service, Edmonton, 
Alberta; personal communication), each needing 40 
seals/yr, their take should be of the order of 0.7x10^ 
seals/yr; i.e., an order of magnitude larger than the 
human harvest. 

The Arctic Fox is frequent on the sea ice in winter. 
It kills newborn pups, digging into the under-snow 
lairs to get them. It has been estimated that locally as 
much as 40% of the annual production of pups may 
be lost to fox predation (Smith 1976), but there is no 
estimate of the average rate of this mortality. 

The fourth cause of natural mortality is intra- 
specific aggression. Sub-adult seals in fast-ice areas 
are liable to be persecuted by adults, which bite them 
on the hind flippers and in the axillae, to a point 
where they are too frightened to go in the water. They 
stay on the ice or in snow lairs, where they freeze, 
starve or get killed by bears (Stirling and McEwan 
1975; Smith and Hammill 1981; Smith 1987). 

Total mortalities for hunted populations in the 
eastern Canadian Arctic were estimated from 
analysis of a catch curve at 23% for pups, 1 2 to 13% 
for sub-adults, and 10 to 1 1% for adults, of which 
natural mortalities were 13%, 5 to 9%, and 3% 
(Smith 1973b). In the eastern Arctic, rates were 16% 
for pups and 14% for other ages (Smith 1987). 

Special Significance of the Species 

The Ringed Seal is still important as food for man 
and dog in the Arctic, now less so as a source of fuel 
and clothing. This species is unique in its special 
association with the Polar Bear. Without the Ringed 
Seal, the Polar Bear could not survive in its present 
numbers. Canada is under international obligation 
to protect Polar Bear populations and their habitat. 
The Ringed Seal is also an important food source 
for Arctic Foxes in some areas and at some times of 
year. Polar Bear and fox offer the Inuit 
opportunities, through trapping and hunting, to use 
their traditional skills and aptitudes to earn cash. 

Evaluation 

There is now no reason to place the Ringed Seal in 
any category under COSEWIC. 

Acknowledgments 

Studies on the Ringed Seal in the Canadian Arctic 
have been supported by the Department of Fisheries 
and Oceans and the Department of Indian and 
Northern Affairs, and information gathered by the 
Canadian Wildlife Service in its studies on the Polar 
Bear has been of great value. These studies have 
been supported in kind by the Polar Continental 
Shelf Project. I thank T. G. Smith and I. Stirling for 



144 



The Canadian Field-Naturalist 



Vol. 104 



sharing their knowledge of this species. Studies on 
the distribution, abundance and behaviour of the 
Ringed Seal have also been supported by the 
Beaufort Sea Project, the Arctic Islands Pipeline 
Project, the Eastern Arctic Marine Environmental 
Studies program, and, in Alaska, the Outer 
Continental Shelf Environmental Assessment 
Program; also by the Arctic Islands Operating 
Advisory Committee, Dome Petroleum Ltd., Esso 
Resources Canada Ltd., Canarctic Shipping Ltd., 
Gulf Canada Resources Inc., Polar Gas Ltd., and 
the Environmental Studies Revolving Funds. I 
thank H. J. Cleator for help in preparing this article. 

Literature Cited 

Alliston, W. G. 1980. The distribution of ringed seals in 
relation to winter icebreaking activities near McKinley 
Bay, N.W.T., January— June 1980. LGL Limited 
Report for Dome Petroleum Limited, Calgary, Alberta. 
52 pages. 

Alliston, W. G., and M. A. MacLaren. 1981. The 
distribution and abundance of ringed seals in western 
Coronation Gulf, Prince Albert Sound, and Minto 
Inlet, N.W.T. LGL Limited Report for Polar Gas 
Project, Toronto, Ontario. 37 pages. 

Burns, J. J., and B. P. Kelly. 1982. Studies of ringed seals 
in the Alaskan Beaufort Sea during winter: impacts of 
seismic exploration. Annual report, 1982, prepared for 
the Outer Continental Shelf Environmental Asessment 
Program, Contract NA 8 1 RAC 45. Alaska Department 
of Fish and Game, Fairbanks, Alaska. 

Burns, J. J., L. H. Shapiro, and F. H. Fay. 1980. The 
relationships of marine mammal distributions, densities 
and activities to sea ice conditions. Final Report, 
OCSEAP contract 03-5-022-55. Alaska Department of 
Fish and Game, Fairbanks, Alaska. 

Calvert, W., and I. Stirling. 1985. Winter distribution of 
ringed seals {Phoca hispida) in the Barrow Strait area, 
Northwest Territories, determined by underwater 
vocalizations. Canadian Journal of Fisheries and 
Aquatic Sciences 42: 1238-1243. 

Canada Department of Fisheries and Oceans. C.R.C. 

1978. Seal Protection Regulations made under the 
Fisheries Act. Published by authority of the Minister, 
Department of Fisheries and Oceans, Ottawa, Ontario. 

Davis, R. A., K. J. Finley, and W. J. Richardson. 1980. 
The present management and future status of Arctic 
marine mammals in Canada. Report prepared for the 
Science Advisory Board of the N.W.T. Department of 
Information, Government of the Northwest Territories. 
93 pages. 

Digby, S. A. 1984. Remote sensing of drained ice areas 
around the breathing holes of ice-inhabiting seals. 
Canadian Journal of Zoology 62: 101 1-1014. 

Eberhardt, L. L. 1978. Transect methods for population 
studies. Journal of Wildlife Management 42: 1-31. 

Eberhardt, L. L., D. G. Chapman, and J. R. Gilbert. 

1979. A review of marine mammal census methods. 
Wildlife Monographs 63. 46 pages. 

Fay, F. H. Carnivorous walrus and some Arctic 

zoonoses. Arctic 13: 111 122. 
Finley, K.J. 1976. Studies of the status of marine 

mammals in the central district of Franklin, N.W.T., 



June-August 1975. LGL Limited Report for Polar Gas, 
Toronto, Ontario. 183 pages. 

Finley, K. J. 1979. Haul-out behaviour and densities of 
ringed seals, {Phoca hispida), in the Barrow Strait area, 
N.W.T. Canadian Journal of Zoology 57: 1985-1997. 

Finley, K. J., and W. G. Johnston. 1977. An investiga- 
tion of the distribution of marine mammals in the 
vicinity of Somerset Island with emphasis on Bellot 
Strait, August-September 1976. LGL Limited Report 
for Polar Gas, Toronto, Ontario. 91 pages. 

Finley, K. J., G. W. Miller, R. A. Davis, and W. R. 
Koski. 1983. A distinctive large breeding population of 
ringed seals {Phoca hispida) inhabiting the Baffin Bay 
pack ice. Arctic 36: 162-173. 

Frost, K. J., and L. F. Lowry. 1981. Ringed, Baikal and 
Caspian seals. Pages 29-53 in Handbook of marine 
mammals. Edited by S. H. Ridgway and R. J. Harrison. 
Academic Press, London. 

Frost, K. J., and L. F. Lowry. 1984. Trophic relation- 
ships of vertebrate consumers in the Alaskan Beaufort 
Sea. Pages 381-401 in The Alaskan Beaufort Sea: eco- 
systems and environments. Academic Press, London. 

Frost, K. J., L. F. Lowry ,and J. J. Burns. 1985. Ringed 
seal monitoring: relationships of distribution, 
abundance, and reproductive success to habitat 
attributes and industrial activities. Annual report, 1985, 
prepared for the Outer Continental Shelf Environmen- 
tal Assessment Program, Contract 84-ABC-00210. 
Alaska Department of Fish and Game. 85 pages. 

Hertz, O., and F. O. Kapel. 1986. Commercial and 
subsistence hunting of marine mammals. Ambio 15: 
144-151. 

King, J. 1983. Seals of the world. British Museum and 
Cornell University Press, Ithaca, New York. 240 pages. 

Kingsley, M. C. S. 1979. Fitting the von Bertalanffy 
growth equation to polar bear age-weight data. 
Canadian Journal of Zoology 57: 1020-1025. 

Kingsley, M. C. S. 1984. The abundance of ringed seals 
in the Beaufort Sea and Amundsen Gulf, 1983. 
Canadian Manuscript Report on Fisheries and Aquatic 
Sciences 1778. 8 pages. 

Kingsley, M. C. S. 1986. Distribution and abundance of 
seals in the Beaufort Sea, Amundsen Gulf, and Prince 
Albert Sound. Environmental Studies Revolving Funds 
Report 25. 16 pages. 

Kingsley, M. C. S. 1989. The distribution of hauled-out 
ringed seals and an interpretation of Taylor's law. 
Oecologia79: 106 110. 

Kingsley, M. C. S., and T. W. Byers. 1988. Status of the 
ringed seal population of Thesiger Bay, N.W.T., 1987. 
Report Prepared for the Fisheries Joint Management 
Committee. Department of Fisheries and Oceans, 
Ottawa, Ontario. 30 pages + tables. 

Kingsley, M. C. S., and T. W. Byers. 1989. Status of the 
ringed seal population in Thesiger Bay, N.W.T., 1987- 
1988. Report Prepared for the Fisheries Joint 
Management Committee. Department of Fisheries and 
Oceans, Ottawa, Ontario. 

Kingsley, M. C. S., L Stirling, and W. Calvert. 1985. The 
distribution and abundance of seals in the Canadian 
high Arctic, 1980-1982. Canadian Journal of Fisheries 
and Aquatic Sciences 42: 1189 1210. 

Kumlien, L. 1879. Contributions to the natural history of 
Arctic America made in connection with the Howgate 
Polar Expedition, 1877-78. Bulletin of the U.S. National 
Museum 15. 179 pages. 



1990 



KiNGSLEY: STATUS OF THE RINGED SEAL 



145 



Lydersen, C, and T. G. Smith. 1989. Avian predation on 
ringed seal, Phoca hispida, pups. Polar Biology 9: 
489-490. 

Lowry, L. F., and F. H. Fay. 1984. Seal eating by wal- 
ruses in the Bering and Chukchi seas. Polar Biology 3: 
11-18. 

Lowry, L. F., K. J. Frost, and J. J. Burns. 1980. Variabil- 
ity in the diet of ringed seals, Phoca hispida, in Alaska. 
Canadian Journal of Fisheries and Aquatic Sciences 37: 
2254-2261. 

McLaren, L A. 1958a. The biology of the ringed seal, 
{Phoca hispida Schreber), in the eastern Canadian 
Arctic. Bulletin of the Fisheries Research Board of 
Canada 118. 97 pages. 

McLaren, L A. 1958b. The economics of seals in the 
eastern Canadian Arctic. Fisheries Research Board of 
Canada, Arctic Unit Circular 1. 94 pp. 

McLaren, LA. 1961. Methods of determining the 
numbers and availability of ringed seals in the eastern 
Canadian Arctic. Arctic 14: 162-175. 

McLaren, L A. 1966. Analysis of an aerial census of 
ringed seals. Journal of the Fisheries Research Board of 
Canada 23: 769-773. 

Smith, T. G. 1973a. Censusing and estimating the size of 
ringed seal populations. Fisheries Research Board of 
Canada Technical Report 427. 18 pages + figures. 

Smith, T. G. 1973b. Population dynamics of the ringed 
seal in the Canadian eastern Arctic. Bulletin of the 
Fisheries Research Board of Canada 181. 55 pages. 

Smith, T. G. 1975. Ringed seals in James Bay and 
Hudson Bay: population estimates and catch statistics. 
Arctic 28: 170-182. 

Smith, T. G. 1976. Predation of ringed seal pups, {Phoca 
hispida) by the Arctic fox {Alopex lagopus). Canadian 
Journal of Zoology 54: 1610-1616. 

Smith, T.G. 1979. How Inuit trapper-hunters make ends 
meet. Canadian Geographer 99(3): 56-61. 

Smith, T. G. 1987. The ringed seal, Phoca hispida, of the 
Canadian western Arctic. Canadian Bulletin of Fisheries 
and Aquatic Sciences 216. 81 pages. 

Smith, T. G., and M. O.Hammill. 1980a. Asurveyofthe 
breeding habitat of ringed seals and a study of their 
behaviour during the spring haul-out period in south- 
eastern Baffin Island. Canadian Manuscript Report on 
Fisheries and Aquatic Sciences 1561. 47 pages. 

Smith, T. G., and M. O. Hammill. 1980b. Ringed Seal, 
Phoca hispida, breeding habitat survey of Bridport Inlet 
and adjacent coastal sea ice. Canadian Manuscript 
Report on Fisheries and Aquatic Sciences 1577. 31 
pages. 

Smith, T. G., and M. O. Hammill. 1981. Ecology of the 
ringed seal, Phoca hispida, in its fast ice breeding 
habitat. Canadian Journal of Zoology 59: 966-981. 

Smith, T. G., and L Stirling. 1975. The breeding habitat 
of the ringed seal {Phoca hispida). The birth lair and 
associated structures. Canadian Journal of Zoology 52: 
1297-1305. 

Smith, T. G., and L Stirling. 1978. Variation in the 
density of ringed seal, {Phoca hispida) birth lairs in the 
Amundsen Gulf, Northwest Territories. Canadian 
Journal of Zoology 56: 1066 1070. 

Smith, T. G., and D. Taylor. 1977. Notes on marine 
mammal, fox and polar bear harvests in the Northwest 
Territories, 1940 to 1972. Environment Canada, 
Fisheries and Marine Service Technical Report 694. 37 
pages. 



Smith, T. G., K. Hay, D. Taylor, and R. Greendale. 

1978. Ringed seal breeding habitat in Viscount 
Melville Sound, Barrow Strait and Peel Sound. 
ESCOM A 1-22. Arctic Islands Pipeline Programme, 
Environmental-Social Committee, Northern Pipeline 
Agency, Ottawa, Ontario. 85 pages. 

Smith, T. G., G. A. Sleno, and D. Taylor. 1979. An 
aerial survey of marine mammals in the region of 
Cornwallis Island, N.W.T. Environment Canada, 
Fisheries and Marine Service Technical Report 837. 14 
pages. 

Stewart, R. E. A., P. Richard, M. C. S. Kingsley, and 
J.J. Houston. 1986. Seals and sealing in Canada's 
northern and Arctic regions. Canadian Technical 
Report on Fisheries and Aquatic Sciences 1463. 31 
pages. 

Stirling, I. 1973. Vocalization in the ringed seal (P/joca 
hispida). Journal of the Fisheries Research Board of 
Canada 30: 1592 1594. 

Stirling, L 1974. Midsummer observations on the 
behaviour of wild polar bears {Ursus maritimus). 
Canadian Journal of Zoology 52: 1 191-1 198. 

Stirling, L 1977. Adaptations of Weddell and ringed 
seal to exploit polar bear fast ice habitat in the presence 
or absence of predators. Pages 741-748 in Adaptations 
within Antarctic ecosystems. Proceedings of the Third 
SCAR Symposium on Antarctic Biology. Edited by 
G. A. Llano. Smithsonian Institute, Washington, D.C. 

Stirling, L 1983. The evolution of mating systems in 
pinnipeds. Pages 489-527 in Advances in the study of 
mammalian behaviour. Edited by J. F. Eisenberg and 
D. G. Kleiman. American Society of Mammalogists 
Special Publication 7. 

Stirling, I. 1985. Research and management of polar 
bears {Ursus maritimus). Polar Record 23: 167-176. 

Stirling, L 1988. Attraction of polar bear Ursus 
maritimus to offshore drilling sites in the eastern 
Beaufort Sea. Polar Record 24: 1-8. 

Stirling, L, and W. R. Archibald. 1977. Aspects of 
predation of seals by polar bears. Journal of the 
Fisheries Research Board of Canada 34: 1 126 1 129. 

Stirling, L, and E. H. McEwan. 1975. The caloric value 
of whole ringed seals {Phoca hispida) in relation to 
polar bear {Ursus maritimus) ecology and hunting 
behaviour. Canadian Journal of Zoology 53: 
1021-1027. 

Stirling, L, W. R. Archibald, and D. P. DeMaster. 1977. 
Distribution and abundance of seals in the eastern 
Beaufort Sea. Journal of the Fisheries Research Board 
of Canada 34: 976 988. 

Stirling, L, H. Cleator, and T. G. Smith. 1981. Marine 
mammals. Pages 44-58 in Polynyas in the Canadian 
Arctic. Edited by I. Stirling and H. Cleator. Canadian 
Wildlife Service Occasional Paper 45. 

Stirling, L, M. C. S. Kingsley, and W. Calvert. 1982- 
. The distribution and abundance of seals in the 
eastern Beaufort Sea, 1974-1979. Canadian Wildlife 
Service Occasional Paper 47. 23 pages. 

Stirling, L, W. Calvert, and H. Cleator. 1983- 
. Underwater vocalizations as a tool for studying the 
distribution and relative abundance of wintering 
pinnipeds in the high Arctic. Arctic 36: 262-274. 

Accepted 10 October 1989 



Book Reviews 



Zoology 

The Collins Guide to the Birds of Britain and Europe with North Africa and the Middle East 



By Hermann Heinzel, Richard Fitter, and John 
Parslow. 1988. (Reissue of 1984 edition). Stephen 
Greene Press, Lexington, Massachusetts. 320 pp., 
illus. U.S. $15.95. 

This is another reprint of a popular field guide 
by a different pubHsher. The copy that I have on 
hand is sUghtly different from the 1972 first edition 
published by Collins which sold for £1.50 at that 
time. The format is the same but the stock is 
thinner which makes the soft cover edition some 3 
mm thinner. The font used is similar but appears to 
be finer in the new reprint and matches that of the 
revised edition (1984). In fact, the whole book is 
almost identical to that edition. The main 
difference is in the reproduction of the colour 
plates which are in general somewhat duller when 
compared to those of the first edition but the 
difference is not as marked when the 1984 edition is 
used as a reference. 

This guide is the most comprehensive for the 
area it is intended for, covering most of the western 
Palearctic Region. All the species are depicted on 



the colour plates, sometimes in several plumages, 
and a range map is provided for nearly all the 
species, along with a concise text which I always 
found accurate and very useful. In addition, the 
range of the species known to breed in Britain is 
also depicted on individual maps collected in an 
appendix. This field guide will be more useful than 
most and appreciated by those who plan to observe 
birds in Europe, the USSR, the Ural Mountains, as 
well as Iceland and Siptzbergen, the Middle East, 
the Canaries, the Azores, and North Africa. It is 
the most comprehensive and reliable guide 
available for those regions at a very accessible 
price. I highly recommend it to anyone planning 
the travel to Europe and surrounding regions and 
watch birds, whether one may be an avid birder or 
a casual observer. 

Henri Ouellet 



Canadian Museum of Nature, P.O. Box 3443, Station D, 
Ottawa, Ontario K1P6P4 



The Collins Guide to the Birds of South-East Asia 



By Ben King, Martin Woodcock, and E. C. Dickinson. 
1988. (Reissue of 1975 edition). Stephen Greene Press, 
Lexington, Massachusetts. 480 pp., illus. U.S. $19.95. 

This comprehensive guide which covers "all the 
1 198 species that are known to have occurred prior 
to 31 May 1971 in Burma, Malaya, Thailand, 
Indochina (Laos, Cambodia and Vietnam), and 
Hong Kong" is a reprint of the original Collins 
edition first published in Great Britain in 1975. The 
present reprint has a different publisher, but the text 
is similar to that of the earlier version and no new 
titles have been added to the bibliography. The 
quality of the printing remains very good and the 
reproduction of the colour plates is excellent. 



Although this book provides information that is 
now outdated in a few instances, it remains the best 
available guide for this ornithologically very 
diversified part of the world. It is unfortunate that 
the publisher did not take this opportunity to update 
the reprint of this important work. Its price is an 
excellent value and I strongly recommend it to 
anyone interested in the fascinating birds of 
southeast Asia. 

Henri Ouellet 

Canadian Museum of Nature, P.O. Box 3443, Station D, 
Ottawa, Ontario K1P6P4 



Birds of the Middle East and North Africa: A Companion Guide 



By P. A. D. Hollom, R. F. Porter, S. Christensen, and Ian 
Willis. 1988. Buteo Books, Vermillion, South Dakota. 
280 pp., illus. + plates. U.S. $32.50. 

This field guide covers an immense sector of the 
F'alearctic region "in North Africa, all the countries 
lying between the Mediterranean and the Sahara, 



and, in South-west Asia, Turkey and Iran, and all 
countries southwards." It is defined as a 
"companion guide" because it is meant to 
complement and provide more information than is 
given in European field guides mainly for the species 
not known to occur regularly in Europe. It differs 



146 



1990 



Book Reviews 



147 



from those guides in that the information is the most 
recent available, that the maps are sufficiently large 
and clear to define quite precisely the range of the 
species in the area described above, and that it 
furnishes the user with colour illustrations for 350 
species seldom illustrated in other reference material 
and over 100 line drawings to illustrate points 
mentioned in the text. 

Each species' entry comprises an extensive section 
entitled "Identification" but for those species not 
treated at length in other companion guides this 
section is quite brief, and includes a useful section on 
voice. The section on "Status" varies greatly in 
length but is usually placed near the range map of 
the species treated. Finally, the short section on 
habitat may contain brief information about 
nesting. Although the maps are clear and the range 
of the species can be "seen" at a glance because it is 
boldly indicated in bright red, the user should read 
carefully the introduction to interpret them 
correctly. The text is concise and accurate at least for 
the twenty or so species that I have checked against 
other descriptions or specimens. The authors often 
call attention to recognizable subspecies of 
European birds which can be of special interest to 



those familiar with birds in Europe but not familiar 
with the taxa covered in the guide. 

The line drawings are particularly pleasant and 
well done; their location in the text should prove to 
be particularly useful to the user in the field. The 40 
colour plates by Ian Willis are very attractive if not 
always perfectly accurate. They should be 
particularly valuable to those not well acquainted 
with the birds of that region. However, I am left 
wondering on how many species one could identify 
correctly using the colour plates under the taxing 
conditions that one is likely to encounter in this part 
of the world! 

To sum up, I have no great hesitation in recom- 
mending this guide to whoever plans to observe 
birds in such a fascinating region. The price is right 
and the book is the best available to this date for the 
region, although one will need to take along one of 
the companion guides, particularly if one is from 
North America and does not know European birds 
too well. 

Henri Ouellet 

Canadian Museum of Nature, P.O. Box 3443, Station D, 
Ottawa, Ontario K1P6P4 



An Annotated Bibliography of the Pike, Esox lucius (Osteichthyes: Salmoniformes) 



By E. J. Grossman and J. M. Casselman. 1987. Life 
Sciences Miscellaneous Publication. Royal Ontario 
Museum, Toronto, Canada, xix + 386 pp. $18. 

This publication provides a functional list of 
references dealing with the biology of a well-known 
fish species; the northern pike or pike (Esox lucius). 
Compiled by two renowned esocid specialists, this 
contribution, which lists over 3000 references, will 
be a very useful bibliographic source for many years 
to come. 

Published bibUographies are time-saving tools. 
With the number of scientific and popular journals 
presently found on the shelves of libraries, one can 
easily feel overwhelmed by the task of doing a 
literature search on a specific subject, specially one 
that has been treated as extensively as Esox lucius.). 

The book begins with an essay entitled "The 
Protohistory of Pike in Western Culture" by 
Richard C. Hoffman. We learn that the first direct 
literary reference to the pike (in fact to lucius) was 
made by a Roman poet, Decimus Magnus 
Ausonius (310-393 A.D.), in a Latin poem entitled 
Mosella. Hoffman reports various other historical 
facts related to the pike and tries to interpret them 
within the framework of the intellectual preoccu- 
pations of each period. The essay is interesting and 
instructive. 



In my opinion, a good bibliography must show 
the two following qualities: (i) it must be 
exhaustive, and (ii) it must have a good subject 
index in order to provide its user with a quick and 
easy access to specific groups of references. 

The bibliographic portion of the text is 
voluminous (325 pages). References are organized 
in standard alphabetic (authors) and chronological 
orders. Since Esox lucius has a circumpolar 
distribution in the temperate waters of the 
northern hemisphere, the authors were careful not 
to restrict themselves to North American 
references and have made a honest effort to include 
contributions from the United Kingdom, Europe, 
and Asia. Titles are indicated in their original 
language with several followed by an English 
translation. My first language being French, I 
found the absence of diacritical marks a bit 
disturbing. Anglers and naturalists will be happy 
to know that several articles from popular 
magazines are listed. 

The access to this huge reference list is facilitated 
by a good subject index which occupies 51 pages. 
There are 37 categories dealing with various 
biological themes. These categories are explained 
in the Introduction to the bibUography section. 
They include subjects such as: "Accounts by 



148 



The Canadian Field-Naturalist 



Vol. 104 



Geographical and Political Areas", "Anatomy, 
Morphology and Histology", "Angling and 
Record Catches", Culture and Artificial Propaga- 
tion", "Life History and Habitats", "Manage- 
ment", "Marking and Tagging", "Physiology and 
Biochemistry", "Popular Accounts", "Reproduc- 
tion" and "Toxicology and Contaminants". In 
addition to this index, descriptive or informative 
annotations were added to some references in the 
bibliography itself. These are useful to the reader 
since they give a general idea of the publication's 
content without having to hunt for it. 



It is by frequent usage that the quaHties and 
shortcomings of a bibliography become apparent. 
However, I believe that this publication satisfies all 
the basic requirements that will make it a useful 
tool for anybody interested in a fish species 
allegorically referred to as the "aquatic wolf" or the 
"tyrant of the waters". 

Francois Chapleau 

Department of Biology, University of Ottawa, Ottawa, 
Ontario KIN 6N5 



Sharks 

Edited by John D. Stevens with illustrations by Tony 
Pyrzakowski. 1987. Facts on File, New York. 240 
pp., illus. U.S. $29.95; $40.95 in Canada. 

Sharks have a fearsome reputation among the 
uncritical consumers of movies and sensational 
newspaper reports, but bee stings and dog bites are 
more fatal to humans than the maligned shark. 
Once past this undeserved reputation, sharks are a 
fascinating group of fishes with a wide variety of 
body forms, behaviours, physiologies, adapta- 
tions, and other features well worthy of yet another 
book about them. 

This particular book has 17 contributors, in 
addition to the consulting editor and illustrator, 
ranging from professional ichthyologists to 
painters, divers, photographers, and science 
writers with several of these skills being found in a 
single individual. Some of the information in the 
text is not merely compiled from other books but is 
based on experience and original research of the 
contributors. The aim of the book is to provide the 
interested general reader with a balanced and 
comprehensive survey of sharks and their biology. 

The book contains 64 pages on shark attacks in 
different parts of the world. Certainly, sharks will 
attack people but it has been estimated that well 
over 50% of attacks are unrelated to feeding, more 
than 75% of victims were struck only once or twice, 
and less than 30% of attacks were fatal. People are 
just not part of the normal diet of sharks, yet the 
fear of being eaten is morbidly fascinating. Other 
sections deal with evolution, kinds of sharks, 



distribution, biology, senses, ecology, behaviour, 
legends, uses, encountering sharks and repelling 
them, and even how to photograph them. The 
book ends with a checklist, a selected and rather 
short bibliography of 77 items including books and 
certain scientific articles, brief biographies of the 
contributors, and an index. 

This is a large format book replete with superb 
colour photographs and paintings on every page. 
Text is therefore limited but a very good read. 
Errors are few and one I noticed was the record of 
27 people who suffered a "mass attack" in 1949 
with 14 deaths near Ahwaz, Iran. These attacks 
actually occurred spread over the period 1941- 
1949, numbered 24 (and possibly 36 or more) with 
16 (or more) deaths. The original data is somewhat 
vague but it is clear that shark attacks in fresh 
waters draining to the head of the Persian Gulf are 
common and ongoing based on recent studies by 
the reviewer for the period 1938-1985. 

Layout is generally excellent. Some topics are 
separated into boxes, further chopping up the text, 
but I did not find this annoying, perhaps because I 
dipped into this book at intervals rather than 
settling to a solid read. The index works well. 

This book is warmly recommended as the best 
available on a well-worn topic. 

Brian W. Coad 

Ichthyology Section, Canadian Museum of Nature, Box 
3443, Station D, Ottawa, Ontario KiP 6P4 



Suivi de la peche sportive dans Ics eaux de la region de Montreal 



Par P. Fournier, M. Beaudoin, and L. Clout- 
ier. 1987. Gouvernement du Quebec, Ministere du 
Loisir, de la Chasse et de la Peche, Direction g^n^rale 
de Montreal, Service de ramcnagement et de 
I'exploitation de la faune. Rapport technique no. 06- 
42. ix + 76 pp., illus. Free. 



Sport fishing is one of the most popular outdoor 
activities in Canada. Being an ichthyologist but 
also an unconditional adept of sport fishing, I was 
looking forward to reading this short report. It 
contains a survey of the fishing activities in the 



1990 



Book Reviews 



149 



waters surrounding the heavily populated 
Montreal region. The study was done from April 
to mid-October 1985 and had as objectives the 
determination of the diversity of fish species 
caught, the size and weight of these fishes, and the 
most exploited areas in the catchment basins 
around Montreal. Particular attention was given 
to lakes St. Louis, St. Fran9ois, des Deux 
Montagnes, and the St. Lawrence River between 
the Lachine Rapids and the Boucherville Islands. 
The collaboration of local outfitters and sport 
fishing clubs was instrumental in the success of the 
survey. 

In all, 7568 individuals with at least one catch in 
their fishing basket were interviewed. They 
captured a total of 41 273 fishes belonging to 27 
species and representing a total weight of 16 450 
kg. Three species accounted for 86% of all fish 
captured: the Yellow Perch (65%), the Northern 
Pike (14%) and the Walleye (7%). In terms of 
weight, the Northern Pike was the most important 
contributor with 38%, followed by the Yellow 
Perch with 23% and the Walleye with 19%. Each 
interviewed angler returned home with an average 
of 5.45 fishes weighing around 2.2 kg. Fishing 
success and the size of fishes captured varied 
seasonally and geographically. 

This report is full of interesting information. 
Several figures summarizing the biological 
characteristics (size, weight, etc.) of the captured 
fish species for each region and maps indicating the 
most visited (best?) area in each basin for the most 
sought after species are included. 

The M.L.C.P. is to be commended for doing and 
pubUshing this study. Its realization must have 



entailed a great expenditure. Follow-up studies 
will have to be made in order to examine the 
evolution of the exploitative trend. However, to be 
useful, this information will have to be integrated 
in a broader management perspective. In the last 
100 years, changes in the environment have been 
far too extensive and rapid for the ecosystem as a 
whole to absorb without deleterious effects, and 
for biologists to even begin to understand. To this 
day, wildlife management policies have centered 
mainly on the control of the human impact on the 
exploited natural resources (i.e. hunting and 
fishing regulations). More often than not, this has 
been done without a sound knowledge of the 
biology of the resources and of their place within 
the ecosystem as a whole. Research priorities must 
be reoriented toward this last endeavor if we want 
to develop more coherent and fruitful management 
policies. 

On a more technical aspect, the publication is in 
the form of a stapled manuscript. The quality of 
the few photographs is below standard but the 
writing style and line diagrams are clear. A few 
misspellings were noted. Also, I fail to see the 
relevancy of pubhshing, even in a technical report, 
photocopies of computer printout showing results 
of statistical analyses. However, the fact that this 
report is free and that it provides a wealth of 
information about sport fishing in the Montreal 
area far outweigh these minor shortcomings. 

Francois Chapleau 

Department of Biology, University of Ottawa, Ottawa, 
Ontario KIN 6N5 



Guide to the Otoliths of the Bony Fishes of the Northeast Atlantic 



ByTero Harkonen. 1986. Danbiu ApS, Hellerup, 
Denmark. 256 pp., illus. U.S. $80.00. 

This book is meant to be a guide for the 
identification of otoliths ("earstones") from fishes of 
northern European shores. Also included are several 
freshwater and anadromous species, about 15% of 
the total. It is not a taxonomic treatment. The 
author's interests are in the feeding habits of seals 
and cormorants, and faecal samples with their hard 
remains were the most convenient means of 
obtaining this information. A total of 103 fish 
species were examined for their otoliths and over a 
third of these are also found in Canadian waters. 

The book comprises an introduction including 
the anatomy, function, composition, and growth 
of otoliths. An explanation is given on how 
otoliths are used in feeding analyses, including 
methods of estimating size of the fish consumed. 



Methods of preserving and treating otoliths are 
outlined. A key to families and species follows. 

The main part of the text is 97 plates facing a text 
description of the species. Each plate may have up 
to six scanning micrographs of fish otoliths 
showing both the left and right sagitta usually in 
decreasing size and with "inside" and "outside" 
views. The text gives Latin, English, German, 
French, Swedish, and Danish names for each 
species, a general description of the otolith (only 
sagittae, the largest of the three otoliths in the fish 
labyrinth, are described), a sentence summarizing 
the most important characters, line drawings of the 
sagitta in lateral and dorsal view to indicate key 
features, and correlations between otolith length 
and body size based usually on a minimum of 30 
specimens. The book concludes with 63 references 
and a short species index. 



150 



The Canadian Field-Naturalist 



Vol. 104 



This book is useful for identification of sagittal 
otoliths where this is meant as a tool for other 
studies such as diet, archaeological investigations, 
and species and size composition analyses. It is not 
an exhaustive treatment of a particular fauna, such 
as that of northeastern Europe, since many otoliths 
are too small or too rare to figure significantly in 
the samples on which the author's primary work is 
based. Nevertheless, there is a stimulus here for 
others to develop a collection of scanning 
micrographs encompassing particular faunas or 
taxa. This would prove most useful for those 
interested in adding characters to systematic 
analyses or to those whose animals consume the 
smaller species of fish. For both identification and 
systematic works, it would have been most useful if a 
few species had been examined in large series 
including both sexes, all sizes, and distributions 
whether horizontal or vertical. Individual variation 
is difficult to estimate from a few micrographs which 
necessarily reflect an ideal state. 

The text has a number of errors in spelling, 
including scientific names, and of arrangement 



although this does not appear excessive. For 
example, the key on page 45 refers the reader to 
Gadidae on page 146 but this page is occupied by 
Cottidae. A glossary of terms would have been 
useful. Colliculum is defined in the text and pops 
up in the keys but is not illustrated or indexed. 

The most important part of this book to the user 
is the keys. Attempts on my inexperienced part to 
key out otoliths in a blind test indicate that this 
requires considerable practice, all the more so if the 
otoliths are broken or worn. Hence, the scanning 
micrographs, though of "perfect" specimens, are 
most valuable as they show structures in an almost 
three dimensional aspect. These micrographs 
make an otherwise highly-specialized book of 
potential use to workers in fields other than the 
author's and the book can be recommended on this 
basis. 

Brian W. Coad 

Ichthyology Section, Canadian Museum of Nature, P.O. 
Box 3443, Station D, Ottawa, Ontario KIP 6P4 



Biology of the Land Crabs 

Edited by Warren W. Burggren and Brian R. McMahon. 
1988. Cambridge University Press, New York. 479 
pp., illus. U.S. $59.50. 

Biology of the Land Crabs is a comprehensive 
evolutionary and comparative overview of the 
current state of knowledge of a diverse group of 
decapod crustaceans. The editors give a very broad 
definition for the land crabs: all anomuran and 
brachyuran crabs that, due to various types of 
adaptation, are active in air. It is clear that some of 
the contributing authors do not agree with the 
definition but all have complied with it. 

Subjects included cover the range of evolution 
and zoogeography, ecology, behaviour, reproduc- 
tion and growth, and the special physiological 
adaptations required for life out of water. 
Activities described include such fascinating topics 
as tree climbing, sound production, migration, 
response to predators, etc. The chapters are well 



coordinated, with cross references to other parts or 
chapters of the book. And although there are many 
areas of overlap, these are of subject matter not of 
focus: thus different aspects of reproduction are 
discussed under ecology, behaviour, and physiol- 
ogy. There is an appendix that briefly outlines the 
natural history of over eighty of the commoner 
terrestrial species, a hst of references, and three 
indices (to authors, scientific names, and subjects). 
I have rarely encountered such a cohesive multi- 
authored work so excellently edited. Its focus on 
the evolution of adaptations to life out of water 
make it of interest to more than just serious 
carcinologists, and 1 would recommend it to all 
who are studying terrestrial adaptation. 

Diana R. Laubitz 

Zoology Division, Canadian Museum of Nature, P.O. 
Box 3443, Station D, Ottawa, Ontario KIP 6P4 



Analyses in Behavioral Ecology: A Manual for Lab and Field 



By Luther Brown and .Icrry F. Downhowcr. 1988. 
Sinauer Associates, Sunderland, Massachusetts, viii + 
194 pp., illus. U.S. $12.95. 

Behavioural ecology is a fast growing area of 
biology, appealing to researchers and students 
alike. For instructors in this subject, a guide to 



feasible projects serving as exercises for students 
would be a boon, and the present manual is 
designed to fill this almost vacant niche. 

Twenty-seven topics are offered under the 
headings of sensory capabilities, feeding patterns, 
spacing patterns, and reproduction. The latter 85 



1990 



Book Reviews 



151 



pages provide a wide-ranging and admirable 
introduction to the statistical analysis of data, 
including visual presentation of data, circular 
statistics, and multidimensional G tests. 

With the avowed hope that the approaches 
advocated should not only expand understanding, 
but also provide enjoyment, the presentation of 
each topic poses questions, outlines methodology 
for collecting relevant observations, indicates an 
interpretation, and includes references to the 
primary literature. A wide diversity of species 
(including insects, birds, rodents, and fish), 
behavioural situations, and analytical methods are 
laudably integrated into the projects, varying in 
logistical simplicity and seasonal availability. The 



adherence to legal and ethical standards is only 
implicit, but the warning of the possibility of 
allergies to bees is explicit. 

Conceptually, the material incorporates some 
advanced issues such as the controversial mating 
advantage of rare males and the relation of sex 
ratios to local mate competition. For the proper 
presentation of these issues, instructors will need to 
do their homework. They will nonetheless 
welcome this book. 

Patrick Colgan 

Department of Biology, Queen's University at Kingston, 
Kingston, Ontario K7L 3N6 



Botany 

The Flora of the Tobermory Islands, Bruce Peninsula National Park 



By J. K. Morton and Joan M. Venn. 1987. Biology 
Series No. 31. University of Waterloo, Waterloo. 92 
pp., illus. $15.00 

John Morton and Joan Venn have produced 
another in a series of floras for the Georgian Bay- 
Lake Huron region of south-central Ontario. This 
volume complements their earlier flora of 
Manitoulin Island and adjacent smaller islands 
(1984), and covers approximately 24 islands north 
of the tip of the Bruce Peninsula, many of which 
are now included within the new Bruce Peninsula 
National Park. 

The total vascular plant flora of these islands 
stands at 542 species. The authors note that the 
species composition on a given island is subject to 
variation, due to factors such as changing water 
levels and movements of dispersal agents like gulls. 
The smaller shoals are particularly susceptible to 
these factors. These and other factors that have 
affected the past and present floristic composition 
of the study area are amply covered in the 
introductory sections of this book (Geology, 
Climate, Past Glacial History, Recent History, 
History of Botanical Exploration). A fairly 
detailed section on Vegetation and Ecology 
outlines the dominant and characteristic vascular 
plants found in the plant communities that have 
developed on these islands. Each of the major 
islands is described briefly, in terms of topography, 
substrates, and major plant communities. In 
general, these introductory sections provide a good 
context for understanding the flora of these 
islands. However, the section on geology is too 
repetitive. For example, the process of differential 
erosion is explained or mentioned four times in the 
span of less than one page. The phytogeographic 



affinities of members of the flora are briefly 
discussed in the introduction, as well. In this 
connection, I should point out a common error 
regarding the floristic affinity of Little Bluestem 
[Andropogon (-Schizachyrium) scoparius\. It is 
usually assumed to be a prairie species, as it is in 
this book, and it is true that this grass occurs in the 
tall-grass prairie of the mid-west. However, it is 
largely eastern in distribution, and is equally, if not 
more, abundant on the coastal plain of eastern 
North America. It can hardly be considered a 
prairie element, unless all coastal plain species are 
also placed in that category. I also consider Wood 
Millet {Milium effusum) to belong to the northern 
mixed forest flora, not to the eastern deciduous 
forest component, as defined by most phytogeo- 
graphers. The floristic affinities of a few other 
species are up for debate. 

A major portion of this book is devoted to an 
annotated listing of the flora. This is the important 
part of the book. Here, information on status and 
habitat are found. There are no real surprises in the 
species Ust, but the data included here will form an 
important database for park planning and future 
surveys of the islands for the purpose of 
monitoring change. The database function of this 
book is greatly enhanced by the tabular listing of 
the plants on each island. It seems odd, however, 
that the tabular listing is not in the same order as 
the main text. The plant families in the main text 
are arranged in the order found in Gray 's Manual 
of Botany, Eighth edition (1950), whereas the 
checklist is largely alphabetical. I differ with an 
occasional synonym or taxonomic concept in this 
treatment, but in all such cases, these are matters of 
taxonomic opinion. 



152 



The Canadian Field-Naturalist 



Vol. 104 



This flora is rounded out by a series of 
photographs (67) by Donald Gunn and John 
Morton. Many of these are of good to excellent 
quality. They represent only a small proportion of 
the species in the flora, but several species that are 
not commonly photographed are included, and of 
course, the photographs provide a colorful 
backdrop for the text. 

There are two aspects of the production of this 
book that require mention. Firstly, the introduc- 
tory sections (and to a lesser extent, the annotated 
flora) contain quite a few typographical errors. 



Secondly, the book is poorly bound. My copy has 
already fallen apart. 

Although the area covered by this flora is small, 
this work still provides a valuable contribution to 
our knowledge of the distribution of species in 
Ontario. Hopefully, it will also provide valuable 
data for the planners and managers of the new 
national park in which these islands are contained. 

William J. Crins 

New York State Museum, Biological Survey, Albany, 
New York 12230 



Lichens of the Ottawa Region (Second Edition) 



By I. M. Brodo. 1988. Special Publication No. 3. Ottawa 
Field-Naturalists' Club, Ottawa. 115 pp., illus. $9.95 + 
$2.50 postage. 

One of the long-standing burdens of amateur 
lichenology in Canada has been a general lack of 
keys to Canadian lichens. Among the first popular 
keys to appear in print in this country were those of 
Dr. Ernie Brodo, published in Trail and Landscape 
by the Ottawa Field-Naturahsts' Club between 1967 
and 1972. Later, in 1981, those keys were 
reassembled to book form as Lichens of the Ottawa 
Region (Syllogeus No. 29 of the National Museum 
of Natural Sciences). 

The first edition of Lichens of the Ottawa Region 
was well received by the naturalist community (see 
for instance James Case's review in The Canadian 
Field-Naturalist 99: 286-287), and the demand for it 
soon exceeded the supply. Now, having been out of 
print for some years, Dr. Brodo's keys are available 
once again. 

In its second edition. Lichens of the Ottawa 
Region has in many respects come out ahead. To 
begin with, the layout is now more attractive and, 
indeed, more "user friendly". Gone are the page- 
width lines of the introductory sections; in their place 
are double columns, much easier to read. Likewise, 
the print is now tighter and less shambling. But best 
of all, the spine-glued binding of the first edition (my 
copy has long since begun to fall apart) has been 
replaced by a plastic coil binding better suited to the 
extensive page-flipping it is "bound" to receive. 

In his preface to the first edition, Dr. Brodo noted 
that a great deal more exploratory work was needed 
in the Ottawa area before the lichen flora could be 
said to have been adequately sampled. Seven years 
later, the second edition now adds no fewer than 28 
new lichen species to the Ottawa flora, bringing the 
regional total to about 400. (Question: if new lichens 
can still be found by the dozen within 50 km of the 
National Museum of Canada, what mouth-watering 
discoveries await the collector in other parts of the 
country?) 



A once-through of the text has turned up no 
obvious typographical errors or other glitches. For 
the most part, I find the word pictures painted by the 
keys to be lucid and easy to visualize. In the keys to 
the crustose species, however, the tattoo of technical 
jargon can at times be a little overwhelming — but 
this merely reflects the intense focus required in order 
to identify them. Fortunately, things are easier with 
the foliose and fruticose lichens; and even where they 
are not, there is always a well-appointed glossary to 
refer to, happily cross-referenced to some 84 
illustrations. 

The taxonomy is up-to-date and liberal. Most 
users will applaud Dr. Brodo's decision to adopt 
Punctelia, Xanthoparmelia, and other recent generic 
segregates of the classic genus Parmelia. This 
practice brings Lichens of the Ottawa Region in line 
with most other modern North American lichen 
references, including Mason Hale's How to Know 
the Lichens. 

My only real criticism of this book has to do with 
its treatment of outdated lichen names, or synonyms. 
Although synonyms are legion in lichenology just 
now (lichen taxonomists have been working 
overtime during the past decade), they appear only in 
the index, where (even worse) they have been cross- 
referenced only in one direction, i.e. from the earlier 
name to the currently accepted one. In my opinion. 
Dr. Brodo would have done better to have at least 
included synonyms in the keys — if only to make life 
easier for those accustomed to the older taxonomies. 

Best served by Lichens of the Ottawa Region are, 
of course, residents of the Ottawa area; it will also, 
however, be found useful for much of Ontario and 
Quebec, and applies with fair regularity even to the 
lichens of the west, especially the macrolichens. 
Clear, well-illustrated, comprehensive, and up-to- 
date, here is a book that has something worthwhile to 
say to Canadian lichen afficionados wherever they 

Trevor Goward 

Edgewood Blue, Box 131, Clearwater, British Columbia 

VOF I NO 



1990 



Book Reviews 



153 



A Second Checklist and Bibliography of the Lichens and Allied Fungi of British Columbia 



By Willa J. Noble, Teuvo Ahti, George F. Otto, and 
Irwin M. Brodo. 1987. Syllogeus 61. National 
Museum of Natural Sciences, Ottawa K1A0M8. 95 
pp. Free. 

In May of 1967, George Otto and Teuvo Ahti 
brought out a preliminary checklist of the lichens 
of British Columbia, hsting some 569 species in 99 
genera. Now, two decades later, a revised checkhst 
has appeared, and here the totals have increased to 
1 013 species (50 new to North America) in no 
fewer than 205 genera. Clearly, lichenology in the 
west has come of age. 

The list is based largely upon herbarium 
specimens, though where no specimens were 
available the authors have made use of literature 
reports instead. Accepted taxa appear in bold face 
and are often accompanied by a brief notation on 
distribution. Doubtful records and synonyms are 
also included, the latter having been cross- 
referenced to the accepted names. The taxonomy is 
refreshingly up-to-date, and includes most of the 
segregate genera formerly included in, for 
example, Lecanora, Lecidea, Parmelia, and 
Physcia. 

Another up-to-date feature of the checklist is its 
inclusion of lichen parasites and other aUied non- 
lichenized fungi. Implicit in this practice is a 
recognition that the lichen life form is not so much 
a discrete biological or taxonomic unit, as a dietary 
preference of various fungi (for algae). Viewed 
from this perspective, lichenology is really just a 
fuzzy-edged subunit of mycology, and it is 
therefore not unreasonable that lichenologists 
should finally begin to study allied fungal groups. 
Twenty-two of these have been included in the 
present checklist, and it can be expected that many 
more will appear in future revisions. 

Following the checklist is a comprehensive 
bibliography consisting of nearly 300 titles. 
Unfortunately, little attempt has been made to tie 
these references to the checklist itself, and so the 
user is seldom able to track down any particular 



entry to its source. A more helpful approach might 
have been to include at least one specimen citation 
or literature reference for each accepted entry. 
Admittedly, some of this information is already 
available in the original checklist, and some is to be 
found in Brodo's Catalogue of Lichens (1981). The 
former reference, however, is now largely out-of- 
date, and the latter has never been published. 

For the most part, the text is admirably free of 
errors, though given the complexity of a work of 
this kind, it is not surprising that a few mistakes 
have inevitably crept in. Putting aside small points 
of punctuation and spelhng, I note the following: 
1) Erioderma sorediatum (p. 38) was never 
published in that genus, but appeared as 
Leioderma sorediatum, correctly listed on p. 48 
(but note that the author citation should be 
Galloway and P. M. Joerg, and not P. M. Joerg 
and Galloway!); 2) Leptogium cyanescens (Ach.) 
Koerber (p. 49) should actually be cited as 
(Rabenh.) Koerber; 3) the inclusion of Leptogium 
rivale tuck. (p. 49) is based on a misidentified 
specimen of Collemafecundum Degel., and should 
be deleted from the list; 4) Neofuscelia loxodes 
(Nyl.) Essl. has been incorrectly listed as 
'' Melanelia" loxodes (Nyl.) Essl. (p. 51); and 5) the 
"sinense" in Parmotrema sinense (Osbeck) Hale & 
Ahti (p. 58) should actually be spelled "chinense". 

The Second Checklist is a major event in 
Canadian lichenology. Its publication is sure to 
inspire a closer look at one of the richer lichen 
floras in North America. Already, I am aware of 
some dozen species which have yet to be recorded 
for British Columbia, and doubtless there are 
hundreds more to come. The authors are to be 
congratulated on bringing out a work which will 
serve as a cornerstone of western lichenological 
research for years to come. 

Trevor Goward 

Edgewood Blue, Box 131, Clearwater, British Columbia 
VOE INO 



Mosses, Lichens and Ferns of Northwest North America 



By Dale H. Vitt, Janet E. Marsh, and Robin B. 
Bovey. 1988. Lone Pine Publishing, Edmonton. 296 
pp., illus. Cloth. 

Every so often, a book appears which promises 
to introduce to the many a field of study which 
formerly has been the private domain of the 
academic few. Mosses, Lichens and Ferns may be 
just such a book. Or then again, it may not be. For 
the reviewer, it is something of a dilemma: on the 



one hand this field guide is at once attractive, 
authoritative, easy to use, and best of all, 
inexpensive; on the other hand, it can sometimes 
be vague, overly technical, poorly illustrated, and, 
worst of all, inaccurate. 

The book opens in the expected way with a 
chapter introducing the reader to the study of 
cryptogams. Included here are a minicourse on the 
use of binomials, an overview of the climatic and 



154 



The Canadian Field-Naturalist 



Vol. 104 



vegetation zones of western North America, and 
various notes on collecting, preserving, studying, 
and photographing the groups in question. 

The body of the book is then given over to the 
species accounts, which are ranged in four 
chapters, including one on liverworts. The species 
coverage is fairly comprehensive, with 243 mosses 
discussed under 165 entries, 246 lichens under 155 
entries, 40 liverworts under 20 entries, and 44 ferns 
under 28 entries. Each chapter opens with a 
succinct, but generally somewhat technical, 
overview, and is followed by a key to the species, 
also rather technical in the cases of the mosses and 
liverworts. 

A nice touch are the habitat symbols appearing 
with each species account. These provide the 
reader with a quick idea of which species to look 
for where, and where to look for which species. 
Accompanying the symbols are range maps (very 
approximate) of the Northwest. 

Closing the book are a bibliography, glossary, 
and index. The bibliography is reasonably 
thorough, and provides a brief summary of 
contents for each of the 60 references listed. The 
glossary is perhaps less helpful, owing to a 
complete absence of diagrams or even any cross 
references to the introductory sections. The index, 
though adequate, might have been made more 
accessible to the novice through the use of symbols 
denoting the group (i.e. mosses, hepatics, lichens 
or ferns) to which each species belongs. 

The cover illustration is attractive, but not 
particularly well conceived. Where one might have 
expected, given the book's title, to find an 
illustration of a moss or lichen, one finds instead a 
photograph of a liverwort, Conocephalum 
conicum. Although liverworts do appear in the 
text, one must ask whether including a liverwort on 
the cover of a book which claims to be about 
mosses, lichens, and ferns is not likely to be a 
source of at least initial confusion for the reader/ 
browser unacquainted with these groups. 

Also rather puzzling is the inclusion of ferns in 
this book. Ferns, after all, are vascular plants and 
as such, already find adequate treatment in the 
standard floras, not to mention various popular 
guides, notably How to Know the Ferns by J. T. 
Mickel, and Pacific Northwest Ferns and their 
Allies by T. M. C. Taylor. Assuming that the 
authors' primary objective was to win wider 
currency for some of the less appreciated 
cryptogams, perhaps the Myxomycetes (slime 
moulds) would have been a more logical choice. 

Accompanying the text are nearly 500 illustra- 
tions, of which more than 400 are in full colour. It 
is these latter which constitute the book's major 
strength, most being vibrant, legible, taxonomi- 
cally revealing, and, not least, aesthetically 



satisfying. The only major exceptions are the dark- 
coloured species, for which the illustrations are at 
times entirely unhelpful. The photo of Melanelia 
exasperatula (p. 217), for example, is exasperatula 
in the extreme, as are the photos of Andreaea 
rupestris (p. 56) and Atrichum undulatum (p. 60). 
All of these might have been in part salvaged 
through the use of detailed line drawings. 

Other illustrations miss the mark in different 
ways. Thus, the photo oi Athyriumfilix-femina(p. 
272) has this forest-loving species growing in a 
boulder bed, while on page 273 the fronds of 
Gymnocarpium dryopteris are shown in a semi- 
dehydrated state quite atypical of their usual 
appearance. Among the bryophytes, several 
photos illustrate only the gametophyte, and thus 
completely ignore the often more diagnostic 
sporophytic characters. The sporophytes of 
Hypnum circinale, for example, readily separate 
this species from the otherwise similar H. 
subimponens (both p. 103), though only in the 
latter species are capsules illustrated. 

It must be pointed out, moreover, that not all of 
the photos have been correctly labelled. For 
example, "Dendroalsia abietina" {p. 113, bottom 
right) should be corrected to Rhytidiadelphus 
loreus, "Cladonia coniocraea" (p. 199) to C. 
sulphurina, "Cladonia scabriuscula" {p. 208) to C. 
amaurocraea, "Hypogymnia enteromorpha" (p. 
225) to H. metaphysodes, "Peltigera rufescens" (p. 
230) to P. ponojensis, and " P. polydactyla"{p. 233) 
to P. neopolydactyla. Although the last of these 
could be said to represent simply a more 
conservative taxonomy on the part of the authors 
(in this case Janet Marsh) than on the part of the 
reviewer. Marsh would have done better to 
illustrate specimens whose morphology was 
central, not peripheral, to the species concept. 
Among the lichens, this last observation applies 
equally to the photos of Cetraria canadensis (p. 
212), Xanthoparmelia taractica (p. 223), and 
Hypogymnia imshaugii (p. 226). 

No less uneven than the illustrations are the 
written species accounts they accompany. These 
vary from rather good in many cases, to 
inexcusably vague in others. Much of the difficulty 
the novice will experience in using the descriptions 
stems from the fact that no attempt has been made 
to standardize them, even within a single group. 
This has made cross-checking for any given 
character most tedious, and in many cases 
impossible. 

A special word of caution applies to the 
bryophytes. Because of their small size and often 
rather bewildering greenness, mosses and 
liverworts are perhaps intrinsically less easy to 
identify than other groups. The user of Mosses, 
Lichens and Ferns should not be discouraged if 



1990 



Book Reviews 



155 



many common bryophytes resist his best efforts 
with a hand lens. Notwithstanding the promises 
implicit in the keys and illustrations, final 
determination will in many cases still have to await 
critical examination under a microscope. 

Evaluating a multiauthored, multidisciplinary 
work such as Mosses, Lichens and Ferns is not an 
easy task, and any summarizing comments are 
unlikely to hold across the board. However, it is 
probably fair to suggest that the many weaknesses 
and oversights noted above are artifacts of a book 
rushed into publication before its time. One can 
only hope that future editions will display a closer 
attention to detail. Only then is this book Ukely to 



become the epoch-making event it was obviously 
intended to be in the first place. 

None of the above remarks, however, seriously 
detract from the fact that Mosses, Lichens and 
Ferns is already a handsome volume. As well as 
being the only book of its kind yet to have 
appeared for northwest North America, it is also, 
in most ways, a job well done. Trim, pocket-sized, 
and sturdily bound, here is a book that will be at 
home both in the field and, for the ardent, on the 
coffee table. Buy it. 

Trevor Coward 

Edgewood Blue, box 131 Clearwater, British Columbia 
VOE INO 



Illustrated Guide to Some Hornworts, Liverworts, and Mosses of Eastern Canada 



by Robert R. Ireland and Gilda Bellio-Trucco. 1987. 
Syllogeus 62. National Museum of Natural Sciences, 
Ottawa. 205 pp., illus. 

Attempts to produce popular guides to 
bryophytes are few. One of the major reasons is the 
importance of microscopic features to obtain 
critical identifications. It is possible, however, to 
determine many common bryophytes utilizing 
features visible to the naked eye and a simple 10- 
20x hand lens. The guide presented by Ireland and 
Bellio-Trucco, with the assistance of Linda M. 
Ley, gives eastern Canada a useful manual to 
determine nearly one-third of the bryophytes 
reported from that region. This area is defined by 
the authors to include southern Manitoba and all 
of the provinces east of that, but excludes many 
species that are common in the northern 
Territories and arctic environments in the eastern 
provinces. 

The guide treats 235 species belonging to 90 
genera of mosses, 43 genera of liverworts, and 2 
genera of hornworts. The authors have chosen very 
carefully to include the bryophytes that are most 
likely to be encountered and that can be 
discriminated effectively using non-microscopic 
features. This means that they treat approximately 
3 1 % of the total flora. The guide is an introduction, 
and a very good one, produced by professionals 
who have comprehensive experience with these 
plants. Added to this, they are sensitive to non- 
professional users of the guide. 

The guide is designed for a beginner with an 
interest in plants. A discussion is provided to 
present aspects of the life cycle, structure, 
collecting methods and preservation, and 
directions concerning identification through use of 
the guide. Ceneral keys lead the user to the major 
groups of bryophytes (hepatics, mosses, liver- 
worts) and further keys for each of these groups 



guide the user to the names of genera and their 
species. 

For most bryophytes, no popular English names 
are available, reflecting the general oversight of 
these plants by naturalists. For each species, the 
dichotomous key leads to a name, and this species 
name provides information concerning habitat, 
and abbreviated geographic distribution. For each 
species, a figure is given to present pertinent detail 
so that the identified specimen can be compared 
with the illustration. The user should be warned 
that bryophytes often include an intermixture of 
different species and even genera, therefore 
caution needs to be exercised. Furthermore, some 
bryophytes are somewhat plastic in their 
morphology, and may show variant forms related 
to the habitat conditions under which they grew. 

A serious amateur will undoubtedly accept Dr. 
Ireland's generous offer to refer puzzling 
specimens to him. If so, a well labelled and 
prepared specimen should be provided. 

The guide should encourage field naturalists to 
learn more about these attractive and intriguing 
plants that show such diversity in form and 
biology. The availability of this guide should 
encourage teachers to introduce these plants to 
students. For field naturalists, it opens up a new 
spectrum of plants of great interest. 

I have discovered that copies of this publication 
are no longer available free of charge. This is more 
a reflection of current governmental policy 
concerning science than museum policy. The 
charge is apparently modest and adds postal 
charges. 

W. B. SCHOFIELD 

Department of Botany, University of British Columbia, 
Vancouver, British Columbia V6T 2BI 



156 



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Vol. 104 



Mushrooms and Other Fungi of the Midcontinental United States 



By D. M. Huffman, L. H. Tiffany, and G. Knaphus. 
1989. Iowa State University Press, Ames, Iowa. 326 
pp., illus. U.S. $19.95. 

Another in a long line of handbooks for the 
beginning student of mushrooms and other large 
fungi. The proliferation of this type of book simply 
reflects the growing popularity of the subject. The 
handbooks specializing in specific regions are 
much needed. 

This book is quite above the average. There are 
about 200 color pictures and all are a very good 
quality. I particularly like the format which has the 
color picture of the mushroom on the page facing 
the description and discussion of it. 

The book has three principal sections. The 
introduction of 27 pages is very well written and 
illustrated to explain various features of 
mushrooms from hallucinogens to spore prints. 
The main text, of about 270 pages, is devoted to 
describing nearly 250 common species of 
mushrooms. Finally, there is a short glossary 
which will be an invaluable help to the beginner in 
understanding many of the characters of 
mushrooms. The primary name used for each 
mushroom is the scientific name which is, by 
tradition, in Latin. Well-known common names 
are also included and happily the authors did not 
propose or dream-up common names for those 
mushrooms which lack them (as some recent field 
guides have done, with the amusing result we have 



a series of "common names" which have never been 
used before, such as "Pumpkin Ringstalk", and 
seem nonsensical). 

As pleased as I am with this manual, there are a 
few minor lapses that I want users to note. The 
species of Hericium on pages 213-215, are 
confused. Figure 137 is H. ramosum but the 
description is of H. coralloides. Figure 138 is H. 
coralloides and Figure 6B is more my idea of H. 
erinaceus. Although the spores of Ganoderma 
applanatum may appear to have "minute spines", 
the ornamentation of the basidiospore walls is due 
to pitting of the thick brown wall. The spore print 
color for Ganoderma lucidum is brown, not white 
as stated. Phellinus igniarius causes a white rot, not 
a brown rot as stated. In my experience, 
Piptoporus betulinus fruits only on dead trees or 
dead branches and stems of live trees, not on live 
trees as stated. The name of one of the authors for 
Polyporus alveolaris is abbreviated Bond., not 
Bord. 

A fine little book which I do not hesitate to 
recommend. Canadians should not by-pass this 
book because of its title, nearly all the species can 
be found in most parts of southern Canada. 

J. GiNNS 

Biosystematics Research Centre, Agriculture Canada, 
Ottawa, Ontario KIA 0C6 



How to Identify Mushrooms to Genus VI: Modern Genera 



By D. L. Largent and T. J. Baroni. 1988. Mad River 
Press, Eureka, California. 277 pp. U.S. $22.95. 

This is the sixth in this interesting series designed 
to facilitate identifications of mushrooms by 
students and serious amateurs. This volume 
contains two complete sets of keys to the genera of 
gilled mushrooms, one emphasizing macroscopic 
features (part III), and the other microscopic 
features (part 1). Boletes and other fleshy fungi 
lacking gills are excluded. An excellent idea was 
the inclusion of additional specialized keys to 
genera in restricted habitats or on unusual 
substrates, i.e. dung, greenhouses, burn sites, 
conifer cones. Sphagnum, other mosses, parasit- 
ized mushrooms, and dunes (part V). There are no 
illustrations, but reference to illustrations in earlier 
volumes is included. 

One is tempted to compare this book with Rolf 
Singer's monographic treatment of mushroom 
genera (fice Tc\/\ev^, Canadian Field Naturalist 102: 
402-403, 1 988); however, they are not written at the 



same level. While this volume will undoubtedly be 
very useful as an intermediate teaching tool 
between the use of popular guides and Singer's 
monograph, there are many areas where improve- 
ments could be made before a second printing. The 
parameters for the scope of the book are not given. 
It appears to encompass most temperate North 
American and European mushroom genera. The 
endemic European genera Geopetalum, Haasiella, 
and Leucocortinarius are included. However, all 
endemic southern hemisphere genera are lacking, 
as are Asian genera such as Lampteromyces, and 
genera reaching both Asia and North America 
from the south, e.g. Descolea and Termitomyces in 
Asia, and Pyrrhoglossum, Ripartitella and 
Smithiomyces in North America. 

Nearly all genera treated in the keys are 
described in part II, but Plicaturopsis in the 
macroscopic based key, part 111, is not treated. For 
most genera, brief descriptions of macroscopic and 
microscopic features are given, along with selected 



1990 



Book Reviews 



157 



references to published species keys and illustra- 
tions. The list of pubUshed keys is not as complete 
or as up to date as it could have been. Examples of 
missing keys are Hygrophorus by Bird and Grund 
{Proceedings of the Nova Scotia Institute of 
Science 29: 1-131, 1979), Hydropus by Singer 
{Flora Neotropica 32, 1982), Phaeocollybia by 
Horak {Sydowia 29: 28-70, 1977), Hohenbuehelia 
and Resupinatus by Thorn {Mycotaxon 25: 
321-453, 1986), and Cortinarius subgenus 
Bulbopodium by Smith {Bulletin of the Torrey 
Botanical Club 69: 44-64, 1942). 

Comments on the status of acceptance of each 
genus and other features are also given. In many 
cases, it is not clear if the authors themselves accept 
certain genera, or whether only one of the authors 
accepts a genus. Generic coverage, either as 
accepted genera or as synonyms, is not complete, 
even for temperate North America. No mention is 
made of several genera present in North America 
proposed within the last 20 years, e.g. Megacolly- 
bia 1972, Cantharocybe 1973, Caulorhiza and 
Gammundia 1979, Resinomycena 1981, Neolenti- 
nus and Ossicaulis 1985, and Mythicomyces and 
Stagnicola 1986, nor of some of the more unusual 
genera such as Campanella, or recently resurrected 
genera e.g. Lentinula and Tapinella. 

Several errors were noted in the keys and text. In 
part I, Phaeogalera, a genus with germ pores, 
cannot be keyed out to the Cortinariaceae, the 
family to which it is assigned. Similarly, 
Pseudobaeospora with its attached lamellae, 
cannot be keyed to the Lepiotaceae. Species of 



Arrhenia which the authors treat as Leptoglossum, 
with subglobose to lacrymoid spores, cannot be 
keyed to either genus. Reference to Watling (1979) 
on page 1 15, evidently refers to Orton and Watling 
(1979), and the two statements regarding generic 
recognition attributed to Watling 1979 are 
contradictory for Pseudocoprinus on that page. 
Also, contrary to statements in the text, some 
Galerina species lack plages, some Marasmiellus 
species form rhizomorphs, and, worldwide, there is 
more than one species of Cyphellostereum. 

Notwithstanding the problems discussed above, 
do I recommend this book to students and serious 
amateurs? The answer is yes. Used as a teaching 
tool with some modifications, it will be very useful 
and it contains a lot of helpful hints and 
information. The multiple keys offer students and 
amateurs different options and even where there 
appears to be a problem in one key, the genus can 
often be reached in another or in the tables given 
on pages 140-153. For those who wish to acquaint 
themselves with many of the modern genera, this 
publication is a bargain compared to purchasing 
more technical manuals, and therefore fills a much 
needed gap in the literature. Final words of advice 
to the authors: move the index to the back of the 
book, index the generic names in the keys, and 
make the book comprehensive for a defined 
geographic area. 

S. A. Redhead 

Biosystematics Research Centre, Agriculture Canada, 
Ottawa, Ontario KIA 0C6 



Environment 

Stones of Silence — Journeys in the Himalaya 

By George B. Schaller. 1988. Originally published by 
Viking Press, 1980. University of Chicago Press, 
Chicago. 292 pp., illus. U.S. $14.95. 

As the director of science for the New York 
Zoological Society, George Schaller has roamed 
the globe studying the ecological and evolutionary 
relationships of some of the world's more 
glamorous larger mammals including gorillas and 
Uons in Africa, tigers in India, pandas in China, 
and wild sheep goats in the Himalaya. The results 
of these studies have been published in such titles 
as The Serengeti Lion which received the National 
Book Award for science in 1972. 

His reprinted book. Stones of Silence, is a 
complementary volume to his previous scientific 
monologue, Mountain Monarchs, which is a 
summary of his research on the ecology and 
behavior of Himalayan wild sheep and goats. 



Mountain Monarchs is written for those who want 
only the "facts". Stones of Silence is a more lyrical 
description of his journeys in Pakistan, India, and 
Nepal while conducting the research which lead to 
the publication of Mountain Monarchs. 

While not a scientific treatise. Stones of Silence, 
is nevertheless full of detailed natural history 
observations as well as descriptions of his 
encounters with the land and its people. It is the 
richness of these descriptions which makes the 
book a first class adventure story as much as a 
valuable contribution to science. 

Unlike most scientists, Schaller is an able writer 
and story teller. Stones of Silence shines with 
evocative images of the landscape and the people. 
For example, when Schaller talks about the 
hugiene habits of Tibetans, instead of saying the 
functional "they were dirty", he writes "Tibetans 



158 



The Canadian Field-Naturalist 



Vol. 104 



do not wash unnecessarily". In another, he relates 
how he had to observe burial in Pakistan with a 
body guard to protect him from outlaws. However, 
he found it distracting to do his work "while 
someone behind me played restlessly with the bolt 
of his rifle." While cautiously working his way 
along an ice covered ledge in a narrow gorge, he 
momentarily forgets the dangers when rounding a 
corner and he observes "... a twenty-foot 
waterfall, a magic falls whose tumbling waters 
have been turned to clear shining glass." 

Yet, the overall message of the book is not about 
the beauty of this mountain region as much as it 
relates in graphic detail the rapid destruction of the 
Himalaya flora and fauna as a result of an 
exponentially expanding human population. The 
title. Stones of Silence, poetically refers to what 
Schaller believes is the fate of the Himalaya. He is 
accutely aware that he is witnessing an ecological 
tragedy — a "great dying" as he calls it — 
happening right before his eyes and how the people 
of the region are as much victims as they are 
perpetrators of this impending disaster. He 
described the plight of a forest guard who was 
supposed to prevent timber harvest and hunting in 
a government preserve. Schaller says he saw the 
"vulnerable look of a quiet despair, the eyes of a 
man whose aspirations have been crushed by the 
world's indifference. He could not support his 
family, he told us, not on the one hundred rupees a 
month the government paid him. To survive he 



had to cut and sell the trees he was supposed to 
protect and he had to act as guide to illegal hunters 
..." Yet, Schaller does not condemn the man. 
Later he says "he was a good guide and an honest 
man, his destiny made tragic by fate." 

In essense. Stones of Silence is a metaphor for 
what is happening to the entire Earth, not just the 
Himalaya as human pressures continue to destroy 
the world's ecosystems and simplify the rich 
diversity of life they support. Schaller feels that 
once the region's diverse and abundant wildlife is 
exterminated and extinct, the magnificent 
mountains will remain, but they will be cold, silent 
places, with the warmth of life extinguished. 

Schaller includes the human element as part of 
this tragedy as well. After seeing some crude rock 
pictures of ibex and wolves near one village where 
human destruction of the mountain flora and 
wildlife was proceeding at a rapid rate, Schaller 
writes "When the huts of Besti have turned to dust 
and the last ibex has vanished into the belly of a 
hunter, these petroglyphs will remain speaking of 
the past in a silent voice to silent hills." 

Schaller's book is an adventure story, a natural 
history story, and a conservation story wrapped up 
in one volume. Stones of Silence is a book worth 
reading. 

George Wuerthner 

P.O. Box 273, Livingston, Montana 59047 



Fire in America: A Cultural History of Wildland and Rural Fire 



By Stephen J. Pyne. 1988. (Reprint of 1982 edition). 
Princeton University Press, Princeton. 655 pp., illus. 
U.S. $14.95. 

Stephen J. Pyne is a former fire fighter turned 
history professor who has taken on the exhaustive 
task of trying to write the definitive work on the 
relationship of humans and wild fires — 
particularly as it applies to North America. Pyne 
largely succeeds. 

His comprehensive book. Fire in America, is 
well researched and documented. Introductory 
chapters on fire behavior and fire ecology give 
readers a rudimentary understanding of how fire 
influences the environment. Separate chapters on 
the regional fire histories of the Pacific Northwest, 
Southwest, Northern Rockies, Southeast, Alaska, 
Lake States, and California are included. 

intermixed with these regional historical reviews 
are chapters which document the use of wildfire by 
North American Indian, early European colonists 
and the pioneers who moved westward into the 
Lake States and on to the grasslands beyond. This 



fire-human relationship was largely one of mutual 
benefit. Fires created environments favorable to 
human hunting, gathering, and agricultural 
practices and as a result, humans purposely set 
fires. 

The attitude of using or co-existing with fire 
which punctured the human-fire relationship for 
centuries began to change to control and 
suppression in the late 1800s. According to Pyne, 
the newly created U.S. Forest Service, established 
in 1905, seized wildfire suppression as a means of 
expanding its power and making itself indispensa- 
ble to society. Under Forest Service guidance, 
coordinated federal fire suppression began and 
Pyne then traces the development of fire 
suppression, lead by the Forest Service, among 
federal and state agencies between the 1920s 
through the 1970s. 

In his final chapters, Pyne outlines how scientific 
research on fire ecology during the 1970s has 
brought human attitudes about wild fire full circle 
and resulted in the adoption of a more tolerant view. 



1990 



Book Reviews 



159 



The standard doctrine that the only good fire was a 
dead fire was largely replaced — at least in the 
scientific community — by a perspective which sees 
fire as an essential ecological component of many 
ecosystems. This change in the scientific community 
paved the way for a change in federal fire policy to 
allow at least some fires to burn unhindered. 

Reading Pyne's book gives one the sense that 
humans coevolved with wildfire, that we are, like the 
Lodgepole Pine, a fire-adapted species. Most 
human development and evolution took place in 
regions where periodic wildfires were the norm. Our 
dependence upon fire-adapted or fire-tolerant plant 
and animal communities caused humans to aid the 
expansion of fire's geographic influence by our 
propensity to set ignitions. Just as fire's natural 
geographic influence expands, so did the natural 
range of the human species. Without fire, would Ice 
Age hunters have ever pushed northward into 
Europe or cross the Bering Land Bridge? Reading 
Pyne, one begins to wonder if, along with the 
development of speech, and weapons, the 
partnership with fire might not also have been a 
major influence upon our present evolutionary 
course on earth. 



Though such questions should be of interest to 
more than just those interested in fire history and 
ecology, this title is not hkely to find its way to the 
best seller list. The book's completeness is also one 
of its major drawbacks. It takes quite a 
commitment of time to read and digest all the 
information presented. In addition, chapter 
organization is somewhat confusing. Each one is 
seemingly separate from the one before and after. 
As a result, there is often repetition of the same 
information in different chapters. However, this 
only becomes readily noticeable if one reads the 
book cover to cover in a single sitting. 

Despite these minor shortcomings, Pyne's Fire 
in America is fascinating reading, not only for its 
completeness but the historical perspective it 
brings to today's land management issues. It 
deserves a reading by anyone interested in wildfire 
and its role, not only in natural history, but in 
human history as well. 

George Wuerthner 

P.O. Box 273, Livingston, Montana 59047 



Symbiosis: An Introduction to Biological Association 



By Vernon Ahmadjian and Surindar Paracer. 1986. 
University Press of New England, Hanover, New 
Hampshire. 212 pp., illus. U.S. $32.50. 

According to the authors, this book is primarily 
an introductory text designed for "students, 
instructors, and research workers who wish to 
learn more about symbiosis". I would argue that 
there is something here also for the naturalist. 
Seldom has the thesis that all living things are 
interconnected been so broadly and so cogently 
documented. 

Symbiosis: An Introduction to Biological 
Associations has obviously been written from a 
conviction that the concept of symbiosis has not 
yet received due recognition as one of the unifying 
principles of biology. According to Ahmadjian 
and Paracer, "some of the greatest events in 
biological history, including the origin of the 
eukaryotic cell, as well as the most devastating 
diseases of mankind, are the results of symbiotic 
relationships". 

The book is divided into thirteen chapters, the 
first of which opens with a most readable 
introduction. In it, the authors defend their 
preference for an inclusive definition of symbiosis, 
arguing that that is how the term was originally 
used by Anton de Bary, the German mycologist 
who coined it in 1879. To de Bary, a symbiotic 



association was not (as it is generally understood 
today) always one in which both partners benefit; 
rather it was simply a long-term living together of 
unlike organisms — without regard for the details 
of the relationship. As thus defined, symbiosis 
encompasses the entire spectrum of interspecific 
associations, from parasitism (one partner benefits 
at the expense of the other), through mutualism 
(both partners benefit), to commensalism (one 
partner benefits while the other is unaffected). 

Such a broad definition allows the authors to 
range freely over every major branch of biology, 
there being no organism which does not enter into a 
symbiotic association of one kind or another. A 
simple listing of the major chapter heads reveals the 
breadth of this book: Viral Symbiotic Associations; 
Bacterial Associations; Symbiosis and the Origin of 
the Eukaryotic Cell; Fungal Associations; Protoc- 
tistan Symbiotic Associations; Helminthic Associa- 
tions; Plant Symbiotic Associations; Behavioural 
and Social Symbioses; and, to round things out, 
Symbiosis and Evolution. 

Each of the above topics is discussed in typical 
textbook fashion, with an emphasis on brevity, but 
usually with no corresponding loss of readabiUty. 
Throughout the book, the chapters follow a 
standard format. After a brief introduction to the 
disciphne in question, the authors present a 



160 



The Canadian Field-Naturalist 



Vol. 104 



biological collage in which various symbiotic 
relationships are illustrated by specific examples. 
Closing each chapter is a summary of the main 
points discussed, followed by a suite of well- 
articulated review questions, and then followed 
again by a two-part bibliogrpahy — the first a Ust of 
recommended readings, and the second a listing of 
the more technical literature upon which the chapter 
was based. 

Scattered throughout the book are a half dozen 
"box essays", in which the authors present, in 
popular fashion, various side themes of symbiosis. If 
these were intended to maintain reader interest, I 
can report that they work very well; perhaps space 
will be found in future editions to incorporate more 
of them. 

One distracting feature of this book is the quite 
uneven levels of presentation. Getting through the 
chapter on viral symbiotic associations, for 
example, will probably be a real challenge for the 
amateur naturalist. On the other hand, the 
introduction to the chapter on fungal associations 
reads almost as a junior high school science text. 
Stylistic discrepancies can also be pointed out, 
though perhaps this is unavoidable in a team-written 
book of this kind. 

Putting aside these small points of criticism, 
Symbiosis: An Introduction to Biological Associa- 
tions is a fascinating read. Because the authors have 



undertaken to paint an enormous canvas, and 
because they paint from a perspective unfamiliar to 
most readers, the pictures they present are sure to 
amaze again and again. Did you know, for example, 
that: 

— A heavy infestation of Giardia (Beaver Fever) 
can result in the production of more than 14 
billion cysts per day? 

— England's White Cliffs of Dover consist entirely 
of the fossil remains of foraminiferan amoebae? 

— Trees infected with invading pathogens react by 
cordoning off the infected areas from the rest of 
the tree? 

— Indian Pipe derives part of its nutrient 
requirements from trees through a sort of fungal 
straw? 

— Three out of every four people on earth are 
infected with internal parasites? 

Now that this book exists, I would not be without 
it. If only as a springboard to the vast and 
fascinating literature which surrounds one of the 
primary unifying principles of life. Symbiosis: An 
Introduction to Biological Associations is a valuable 
addition to any naturalist's library. 

Trevor Coward 

Edgewood Blue, Box 131, Clearwater, British Columbia 
VOE INO 



Population Ecology of Individuals 

By Adam Lomnicki. 1988. Monographs of Population 
Biology, 26. Princeton University Press, Princeton, New 
Jersey. 223 p., illus. Cloth U.S. $52; paper U.S. $16. 

Most classic population biology models assume 
no variation between individuals of a population. 
The population is assumed to be composed of 
homogeneous units. Lomnicki challenges this 
assumption. Then, he demonstrates how variation 
between individuals can lead to population stability 
and ultimately ecosystem stability. 

Using graphs and mathematical models to 
illustrate his points, Lomnicki shows how 
population stability could result from two 
mechanisms: unequal resource partitioning between 
individuals and contest competition among 
individuals. Unequal resource partitioning results 
from phenotypic variations between individuals that 
cause differences in resource acquisition abilities. 
Differences in these abilities result in contest 
competition if the resource intake of the larger 
individuals is not affected by the resource intake of 
the smaller individuals. The example of contest 
competition given in this book was competition 



between trees for light. The tallest trees in a forest 
are able to acquire light without interference from 
the shortest trees; however, the light interception by 
the tallest trees reduces the quantity of light acquired 
by the shortest trees. 

Obviously, the quantity of resources that an 
individual acquires is a function of its rank within a 
population. Individuals of the highest ranks often 
acquire resources in greater quantities than the 
resource levels required for reproduction and 
maintenance. Individuals of lower ranks may 
acquire resources between the resource thresholds 
for maintenance and reproduction or even below the 
threshold for maintenance. Individuals acquiring 
resources below the reproduction threshold will not 
reproduce, and individuals acquiring resources 
below the maintenance level will not survive long if 
they remain in the population. Often, the only 
alternative for these last two groups is to emigrate. 

Lomnicki shows how emigration could cause self 
regulation of a population. He also shows how 
perception of a low probability of successful 
reproduction by low ranking individuals could force 



1990 



Book Reviews 



161 



their emigration out of a population in order to 
avoid certain reproductive failure in spite of the 
often low probabilities of finding enough unclaimed 
resources in other populations to meet their 
reproduction requirements. This form of self 
regulation often leaves the population at a level 
lower than the maximum number of individuals that 
its habitat could support. 

This book will appeal to all advanced students, 
scientists, and professionals interested in population 
dynamics. The mathematical models presented in 
this book are relatively simple to follow. Only a 



rudimentary understanding of calculus is required 
to understand these models. Lomnicki uses these 
models to explore some new areas of theoretical 
cology where empirical evidence is not always 
available. Although Lomnicki has done his best to 
use empirical examples, this book's content is 
largely theoretical. However, this author's writing 
style makes this book very accessible to a wide 
audience of biologists. 

Blaine H. M. Mooers 

141 E. Miller, Apt. #6, Sidney, Montana 59270 



Wetlands of Canada 

By the National Wetlands Working Group, Canada 
Committee on Ecological Classification. 1988. 
Environment Canada (distributed by Polyscience, 
Montreal), xiv + 452 pp., illus. $56.00. 

This is a most attractive book. It has an 
abundance of black-and-white photographs and 
well-drawn diagrams. Most of the authors write 
clearly. Everybody interested in this country's 
wetlands will find the chapter dealing with their own 
region absorbing. Therefore, the book deserves the 
attention of nearly all serious amateur naturalists, 
since all of them (surely?) find wetlands fascinating; 
but probably few will want to buy so hefty a book, 
covering the wetlands of all Canada. 

It is the work of 33 contributors, whose 
affihations are: 25 from assorted federal and 
provincial Government departments; 4 from 
ecological consulting firms; and 4 from universities. 
Between them, they have written 10 chapters and an 
appendix. 

The appendix should be read first. It is by a 
committee of 13 of the contributors who present a 
three-level taxonomy of wetlands. At the highest 
level are Classes, of which there are 5 (bogs, fens, 
swamps, marshes, and open water). Next come 
Forms, for a total (in all classes) of 70. Lastly come 
Types, of which there are 16 (on two hierarchical 
levels). None of the forms is represented by more 
than a few possible types, so (fortunately) there are 
fewer than 1 120 ultimate taxa. 

After an introductory chapter. Chapters 2 to 9 
give descriptions of the various wetland taxa in each 
of the 8 Wetland Regions (all with subregions) into 
which Canada is divided. These descriptions are all 
examples (admittedly, good ones) of classical 
descriptive ecology of the kind that prevailed 40 or 
50 years ago. Most are a pleasure to read, and they 



impart plenty of information. But it is rather hke 
listening to a running commentary on the Stanley 
Cup Final on radio instead of switching on the TV. 
The descriptions of vegetation must be based on 
data sets that cry out for modern methods of 
multivariate data analysis, that is, for well-chosen 
classifications and ordinations, but there isn't a sign 
of them. Three (only) of the chapters have diagrams 
showing the putative successional relationships 
among the different wetland taxa in a region. How 
interesting it would be to see ordinations of the data 
on which these diagrams are based! It is not as if 
such a treatment would be too technical for the 
intended readership: the book abounds with 
exceedingly detailed tables giving the results of 
chemical analyses of peat and water samples from 
particular wetlands. 

The indexes are inadequate. There should have 
been an index to plant species and another to 
geographic locations, as well as the rather sketchy 
general index. 

There is a glossary, and it cites the sources of its 
definitions. Perhaps it is unfair to quote the single 
embarrassing howler: "AUTOTROPHIC: Capable 
of deriving energy for life processes from the 
oxidation of inorganic materials (Agriculture 
Canada 1976)." Agriculture Canada should be 
ashamed. 

It is fair to end on a positive note, however. The 
book has many virtues and its faults can be put right 
in promised future editions. One obvious way to 
improve it would be to invite more contributions 
from research workers in the academic world. 

E. C. PlELOU 
RRl, Denman Island, British Columbia VOR ITO 



162 



The Canadian Field-Naturalist 



Vol. 104 



Miscellaneous 



The Outermost House 

By Henry Beston. 1988. (Reprint of 1928 edition). 
Penguin, New York, xxiv + 222 pp. U.S. $6.95; $9.95 in 
Canada. 

The Outermost House, Henry Beston's account 
of a year at Cape Cod, is the latest instalment in the 
Penguin Nature Library series. An instant success at 
the time of its 1928 publication, the book went 
through several reprintings and has now been issued 
in paperback with an introduction by Robert Finch. 

The Finch introduction provides information 
about Beston's life and career, explains the 
circumstances behind the writing of the book, and 
discusses the style and literary merits of the text. The 
reader learns that Beston originally built his cottage 
at Cape Cod, named Fo 'castle, strictly as a summer 
retreat but decided to stay on over the winter of 
1926-27 because he believed that the atmosphere 
would help his writing. The result of his enforced 
exile were several notebooks filled with observations 
and thoughts, which were then transformed, at the 
insistence of his fiancee, into The Outermost House. 

Although Beston liked to describe himself as a 
writer-naturahst. Finch's introduction suggests that 



The Outermost House was largely an exercise in 
creative writing rather than the musings of a 
naturalist. At the same time. Finch notes that Beston 
wonderfully captures the rhythmic quality of life at 
Cape Cod, including the seasonal rituals. 
Unfortunately, Finch does not take his analysis far 
enough to explain why The Outermost House 
became an inspirational piece for the conservation 
movement of the 1960s. Perhaps, this is best left to 
Beston himself who in the powerful closing pages of 
the book warns: "Whatever attitude to human 
existence you fashion for yourself, know it is valid 
only if it be the shadow of an attitude to Nature. A 
human life, so often Ukened to be a spectacle upon a 
stage, is more justly a ritual ... do no dishonour to 
the earth lest you dishonour the spirit of man . . . 
For the gifts of life are the earth's and they are given 
to all ..." (pages 222-223). 

W. A. Waiser 



Department of History, University of Saskatchewan, 
Saskatoon, Saskatchewan S7N OWO 



New Titles 



Zoology 

Advances in comparative and environmental physiology, 
volume 4: animal adaptation to cold. 1 989. Edited by R. 
Gilles, C. P. Mangum, G. N. Somero, K. Takahashi, and 
R. E. Weber. Springer- Verlag, New York. c450 pp., illus. 
U.S.$I13. 

t Alaska whales and whaling. 1988. By Alaska Geogra- 
phic, Anchorage. Reprint of 1978 issue. 144 pp., illus. + 
poster. U.S. $19.95; $23.35 in Canada. 

American warblers: an ecological and behavioral 
perspective. 1989. By Douglas H. Morse. Harvard 
University Press, Cambridge. 384 pp., illus. cU.S.$30. 

The application of remote sensing technology to marine 
fisheries: an introductory manual. 1988. By M. J. A. 
Butler. FAO, Rome, xviii + 165 pp., illus. U.S.$18. 

Arabian mammals: a natural history. 1989. By 
Jonathan Kingdon. Academic Press, San Diego. 300 pp. 
cU.S.$130. 

'''Attracting backyard wildlife: a guide for nature- 
lovers. 1989. By Bill Merilces. Whitecap Books. 
Vancouver, xvi + 159 pp., illus. $12.95. 



tXhe bamboo bears: the life and troubled times of the 
giant panda. 1989. By Clive Roots. Hyperion Press, 
Winnipeg, x + 102 pp., illus. $23.95. 

*Bear-people conflicts. 1989. Edited by Marianne 
Bromley. Proceedings of a conference, Yellowknife, 6- 
10 April 1987. Northwest Territories Department of 
Renewable Resources, Yellowknife. 246 pp., illus. 

*A bibliography of British Columbia ornithology, 
volume 2. 1988. By R. Wayne Campbell, Tracey D. 
Hooper, and Neil K. Dawe. Heritage Record No. 19, 
Royal British Columbia Museum. Crown Publications, 
Victoria. 591 pp. $30. 

*Birds in Ireland. 1989. By Clive B. Hutchinson. Irish 
Wildlife Conservancy (distributed by Buteo, Vermil- 
lion, South Dakota). 215 pp., illus. U.S. $55. 

*The butterflies of Manitoba. 1989. By Paul Klassen, 
Richard Westwood, Bill Preston, and Brian McKillop. 
Manitoba Museum of Man and Nature, Winnipeg, vi + 
290 pp., illus. $21.95 plus $2 postage. 



*Audubon wildlife report 1989/1990. 1989. Edited by 
William .1. Chandler. Academic Press, San Diego. c540 
pp. cU.S.$39.95. 



A dictionary of ethology. 1989. By Klaus Immel Mann 
and Colin Beer. Harvard University Press, Cambridge, 
iii + 335 pp., illus. U.S. $35. 



1990 



Book Reviews 



163 



^Dispersal in rodents: a resident fitness hypothesis. 1989. 
By Paul K. Anderson. Special Publication No. 9, 
American Society of Mammalogists, c/o H. Duane 
Smith, Brigham Young University, Provo, Utah, viii + 
141 pp. 

"'Eastern/central birding by ear: a guide to bird song 
identification. 1989. By Richard K. Walton and Robert 
W. Lawson. Peterson Field Guides. Houghton Mifflin 
(distributed by Thomas Allen, Markham, Ontario). 64 
pp., illus. + 3 tapes. 

Ecological and behavioral methods for the study of bats. 

1988. Edited by Thomas H. Kunz. Smithsonian Institute 
Press, Washington, xxii + 533 pp., illus. U.S. $50. 

Endangered animals. 1988. By Malcolm Penny. 
Bookwright, New York. 32 pp., illus. U.S.S 11.90. 

*Fishes of the Thunder Bay area of Ontario: a guide for 
identifying and locating the local fish fauna. 1989. By 
Connie Hartviksen and Walter Momot. Wildwood Press, 
Thunder Bay. Illus. $24.95. 

Fish to reptiles. 1988. By Lionel Binder. Gloucester, 
New York. 36 pp., illus. U.S.$1 1.40. 

Frozen fauna of the mammoth steppe: the story of Blue 

Babe. 1989. By R. Dale Guthrie. University of Chicago 
Press, Chicago. c360 pp., illus. Cloth cU.S.$40; paper 
cU.S.$16.95. 

Guide to the mammals of Salta Province, Argentina. 

1989. By Michael A. Mares, Ricardo A. Ojeda, and 
Ruben M. Barquez. University of Oklahoma Press, 
Norman. 320 pp., illus. U.S.$29.50. 

Introduction to forest and shade tree insects. 1989. By 
Pedro Barbosa and Michael R. Wagner. Academic Press, 
San Diego. 637 pp. U.S.$59.95. 

The loon: voice to the wilderness. 1988. By Charlene W. 
BilHngs. Dodd, Mead, New York. 48 pp., illus. 
U.S.$11.95. 

tMammals of Oklahoma. 1989. By William Caire, Jack 
D. Tyler, Bryan P. Glass, and Michael A. Mares. 
University of Oklahoma Press, Norman. 544 pp., illus. 
U.S.$29.95. 

*The nature of birds. 1988. By Adrian Forsyth. Camden 
House, Camden East, Ontario. 160 pp., illus. $19.95. 

The new dinosaurs: an alternative evolution. 1988. By 
Dougal Dixon. Salem House, Topsfield, Massachusetts. 
120 pp., illus. U.S.S19.95. 

tOklahoma herpetology: an annotated bibiography. 

1989. By Charles C. Carpenter and James J. Krupa. 
University of Oklahoma Press, Norman. 272 pp. 
U.S.$21.95. 

*0n the track of ice age mammals. 1986. By Anthony J. 
Sutcliffe. Harvard University Press, Cambridge. 224 pp., 
illus. Cloth U.S.$25; paper U.S.$12.95. 



Orang-utan biology. 1988. By Jeffrey H. Schwartz. 
Oxford University Press, New York, x + 383 pp., illus. 
U.S.$79.95. 

tThe reintroduction of the white-tailed sea eagle to 
Scotland: 1975-1987. 1988. By John A. Love. Nature 
Conservancy Council, Peterborough, Great Britain. 48 
pp., illus. 

Shallow-water hydroids of Bermuda: the Athecatae. 

1988. By Dale R. Calder. Royal Ontario Museum, 
Toronto. 107 pp., illus. $24.50. 

The short-tailed fruit bat: a study in plant-animal 
interactions. 1988. By Theodore H. Fleming. University 
of Chicago Press, Chicago, xvi + 365 pp., illus. 
U.S.$49.95. 

*Snakes of eastern North America. 1989. By Carl H. 
Ernst and Roger W. Barbour. George Mason University 
Press (distributed by University Publishing Associates, 
Lanham, Maryland). 282 pp., illus. + plates. U.S. $62.50. 

Snakes of the world, volume 1: synopsis of snake generic 
names; and volume 2: synopsis of living and extinct 
species. 1989. By Kenneth L. Williams and Van 
Wallach. Krieger, Melbourne, Florida. Two volume set 
U.S.$49.50. 

The structure of the call note system of the warbling vireo. 

1989. By Daryl Howes-Jones and Jon C. Barlow. Life 
Sciences Contributions 151. Royal Ontario Museum, 
Toronto. 40 pp., illus. $10.25. 

* Studies of the effects of acidification on aquatic wildlife in 
Canada: lacustrine birds and their habitats in Quebec. 

1989. Edited by Jean-Luc DesGranges. Canadian 
Wildlife Service Occasional Paper No. 67. Supply and 
Services Canada, Ottawa. 70 pp., illus. 

♦Turtles of the world. 1989. By Carl H. Ernst and Roger 
W. Barbour. Smithsonian Institute Press, Washington. 
290 pp., illus. U.S.$45. 

Whale nation. 1988. By Heathcote Williams. Crown, 
New York. 191 pp., illus. U.S.$25. 

t Wildlife and man in Texas: environmental change and 
conservation. 1989. By Robin W. Doughty. Texas A 
and M University Press, College Station. 268 pp., illus. 
U.S.$12.95. 

Botany 

Aerobic photosynthetic bacteria. 1989. Edited by K. 
Harashima, T. Shiba, and N. Murta. Springer-Verlag, 
New York. c200 pp. cU.S.$56. 

Aspects of floral development. 1988. Edited by Peter 
Leins, Shirley C. Tucker, and Peter K. Endress. J. 
Cramer in der Gebruder Borntraeger, Stuttgart, vii + 239 
pp., illus. DM120. 

Atlas florae Europaeae, volume 3: distribution of 
vascular plants in Europe: Caryophyllaceae. 1989. 
Edited by J. Jalas and J. Suominen. Cambridge 
University Press, New York. 405 pp. U.S.$89.50. 



164 



The Canadian Field-Naturalist 



Vol. 104 



The biology and utilization of shrubs. 1988. Edited by 
Cyrus M. McKell. Academic Press, San Diego. 656 pp. 
U.S.$125. 



Plant alert/ Alerte auz plantes. 1990. By Deborah A. 
Metsger. Royal Ontario Museum, Toronto. 64 pp., illus. 
$4.95. 



The biology of polar bryophytes and lichens. 1988. By 
R. E. Longton. Cambridge University Press, New York. 
viii + 391 pp., illus. U.S.$95. 

The conspectus of bryological taxonomic literature, Part 
1: index to monographs and regional reviews; and part 2: 
guide to national and regional literature. 1988 and 1989. 
By S. W. Greene and A. J. Harrington. J. Cramer in der 
Gebruder Borntraeger Verlag, Stuttgart. 272 pp. and 322 
pp. DM 120 each. 

Diatoms. 1989. By F. E. Round, R. M. Crawford, and 
D. G. Mann. Cambridge University Press, New York. 
c550 pp., illus cU.S.$150. 

t Discovering wild plants: Alaska, western Canada, the 
northwest. 1989. By Janice J. Schofield. Alaska 
Northwest, Edmonds, Washington. c350 pp., illus. 
U.S.$34.95; $43.95 in Canada. 

fThe ecology of intercropping. 1989. By John Vander- 
meer. Cambridge University Press, New York, xi + 237 
pp., illus. U.S.$59.50. 

The ecology of soil seed banks. 1989. Edited by Mary A. 
Leek, V. Thomas Parker, and Robert L. Simpson. 
Academic Press, San Diego. 444 pp. U.S. $69. 95. 

Environmental stress in plants. 1989. Edited by J. H. 
Cherry. Springer- Verlag, New York, viii + 369 pp., illus. 
cU.S.$107.50. 

*The genus Vacciniutn in North America. 1989. By S. P. 
Vander Kloet. Agriculture Canada. Canadian Govern- 
ment Publishing Centre, Ottawa. 203 pp., illus. $46.50 in 
Canada; U.S. $55. 80 elsewhere plus $2.25 shipping. 

Grass: systematics and evolution. 1988. Edited by 
Thomas R. Soderstrom, et al. Smithsonian Institute 
Press, Washington, xiv + 473 pp., illus. U.S. $45. 

The Hepaticae of southern Greenland. 1989. By Rudolf 
M. Schuster. J. Cramer in der Gebruder Borntraeger 
Verlag, Stuttgart. 254 pp., illus. DM 170. 

* Indicator plants of coastal British Columbia. 1989. By 
A. Kiinka, V. J. Krajina, A. Creska, and A. M. Scagel. 
University of British Columbia Press, Vancouver, ix + 288 
pp., illus. $36.95. 

Leaf venation patterns, volume 3: Myrtaceae. 1988. By 
Edward P. Klucking. J. Cramer in der Gebruder 
Borntraeger Verlag, Stuttgart. 279 pp., illus. DM 260. 

Modern methods in orchid conservation. 1989. Edited 
by H. W. Prilchard. Cambridge University Press, New 
York. c200 pp. cU.S.$49.50. 

Morphology of flowers and inflorescences. 1989. By F, 
Wcberling. Translated by R. J. Pankhurst. Cambridge 
University Press, New York. c4!6 pp. cU.S.$l 10. 



* Plants of Riding Mountain National Park, Manitoba. 

1989. By William J. Cody. Agriculture Canada. 
Canadian Government Publishing Centre, Ottawa. 319 
pp., illus. $24.75 in Canada; U.S. $29. 70 elsewhere plus 
$1.90 shipping. 

Progress in botany, volume 50. 1989. Edited by H. D. 
Behnke, K. Esser, K. Kubitzki, M. Runge, and H. Ziegler. 
Springer-Verlag, New York. c400 pp., illus. 
cU.S.$147.50. 

Tropical forests and botanical diversity. 1 989. Edited by 
L. B. Holm-Nielson, H. Balslev, and I. Nelson. Academic 
Press (Harcourt Brace Jovanovich, San Diego). 400 pp. 
cU.S.$49.50. 

Vascular epiphytes: general biology and related biota. 

1989. By David H. Benzing. Cambridge University 
Press, New York. 320 pp., illus. cU.S. $42.50. 

Vegetation mapping. 1988. Edited by A. W. Kuchler 
and J. S. Zooneveld. Kluwer, Norwell, Massachusetts, x 
+ 635 pp. U.S.$250. 

Where the gods reign: plants and peoples of the 
Colombian Amazon. 1988. by Richard Evans Schultes. 
Synergetic Press, Oracle, Arizona. 306 pp., illus. U.S. $20. 

Wildflowers of the southern interior of British Columbia 
and adjacent parts of Washington, Idaho, and Montana. 

1989. By Joan Burbridge. University of British 
Columbia Press, Vancouver. c400 pp., illus. Cloth $29.95; 
paper $19.95. 

*Wild rice in Canada. 1988. By S. G. Aiken, P. F. Lee, 
D. Punter, and J. M. Stewart. Agriculture Canada 
(distributed by NC Press, Toronto). 130 pp., illus. $18.95 
in Canada; U.S. $22. 75 elsewhere. 

Woody plants: evolution and distribution since the 
tertiary. 1989. Edited by F. Ehrendorfer. Proceedings of 
a symposium Halle/ Saar, Germany, 9-1 1 October, 1986. 
Springer-Verlag, New York, v + 329 pp., illus. cU.S.$177. 

Environment 

Acidic deposition and forest soils. 1989. By D. Binkley, 
C. T. Driscoll, H. L. Allen, P. Schoeneberger, and D. 
McAvoy. Springer-Verlag, New York, viii + 149 pp., illus. 
U.S.$49. 

Aquatic toxicology and environmental fate, volume 11. 

1989. Edited by Glen W. Suter and Michael A. Lewis. 
ASTM, Philadelphia. 616 pp., illus. U.S. $74. 

jAt the water's edge: nature study in lakes, streams, and 
ponds. 1989. By Alan M. Dvancara. Wiley, Somerset, 
New Jersey, vii + 232 pp., illus. U.S. $10.95. 

Between two worlds: science, the environmental 
movement, and policy choice. 1989. By Lynton K. 
Caldwell. Cambridge University Press, New York. c200 
pp. cU.S.$39.50. 



1990 



Book Reviews 



165 



Biodiversity. 1988. Edited by E. W. Wilson. Based on a 
forum, Washington, September, 1986. National 
Academy Press, Washington, xiv + 521 pp., illus. Cloth 
U.S.$32.50; paper U.S.$19.50. 

Biospheres: from the earth to space. 1988. By Dorion 
Sagan and Lynn Margulis. Enslow, Hillside, New Jersey. 
96 pp. U.S.$14.95. 

The Boundary Waters Canoe Area: wilderness values and 
motorized recreation. 1989. By James N. Gladden. 
Iowa State University Press, Ames. 168 pp., illus. 
cU.S.$18.95. 

Canadian sources of environmental information 1988. 

1989. Canadian Government Publishing Centre, Ottawa. 
457 pp. $39.75 in Canada; U.S.$47.50 elsewhere. 

Conservation biology in Hawaii. 1988. Edited by 
Charles P. Stone and Danielle B. Stone. University of 
Hawaii Press, Honolulu, xxiv + 410 pp., illus. U.S. $16. 

The earth's fragile systems: perspectives on global change. 

1988. Edited by Throkil Kristensen and Johan Peter 
Paludan. Westview, Boulder, Colorado, xiii + 109 pp., 
illus. U.S.$30. 

Ecological relationships of plants and animals. 1988. By 
Henry F. Howe and Lynn C. Westley. Oxford University 
Press, New York, xiii + 273 pp., illus. U.S.$29.95. 

Environmental management handbook: toxic chemical 
materials and wastes. 1989. Dekker, New York, xvi + 
632 pp. U.S.$125. 

Functional testing of aquatic biota for estimating hazards 
of chemicals. 1989. By John Cairns, Jr. and James R. 
Pratt. ASTM, Philadelphia. 242 pp., illus. U.S.$45. 

The future of the environment. 1988. Edited by David 
C. Pitt. Routledge, Chapman, and Hall, New York. 218 
pp. U.S.$49.95. 



Novel aspects of insect-plant interactions. 1988. Edited 
by Pedro Barbosa and Deborah K. Letourneau. Wiley- 
Interscience, New York, xx + 362 pp., illus. U.S.$47.50. 

Our natural resources and their conservation. 1988. By 
Harry B. Kircher, Donald L. Wallace, and Dorothy J. 
Gore. Interstate, Danville, Illinois, xxiii + 482 pp., illus. 
U.S.$19.95. 

Range development and improvements. 1989. By John 
F. Vallentine. Third edition. Academic Press, San Diego. 
524 pp. U.S.$45. 

Rehabilitating damaged ecosystems. 1988. Edited by 
John Cairns. Two volumes. 192 pp. and 224 pp. U.S.$1 10 
each in U.S.A.; U.S.$125 each elsewhere. 

The rights of nature: a history of environmental ethics. 

1989. By Roderick Frazier Nash. University of 
Wisconsin Press, Madison, xiv + 290 pp. U.S.$27.50. 

Valley of the cranes: exploring Colorado's San Luis 
Valley. 1988. By Virginia McConnnell Simmons. 
Roberts Rinehart, Boulder, Colorado. 64 pp., illus. 
U.S.$12.95. 

* Walking the wetlands: a hiker's guide to common plants 
and animals of marshes, bogs, and swamps. 1987. By 
Janet Lyons and Sandra Jordon. Wiley, Somerset, New 
Jersey, xviii + 222 pp., illus. U.S. $10.95. 

fWild Britain: a traveller's and naturalist's guide. 1989. 
By Douglas Botting. Prentice Hall, New York. 224 pp., 
illus. U.S.$13.95. 

Wild Spain. 1989. By Frederick V. Grunfeld. Prentice 
Hall, New York, illus. U.S.$13.95. 

Miscellaneous 

Advances in marine biology, volume 25. 1989. Edited 
by J. H. S. Blaxter and A. J. Southward. Academic Press 
(Harcourt Brace Jovanovich, San Diego). c288 pp. 
cU.S.$70. 



♦Guide to the Queen Charlottes, 1989-1990. 1989. By 
Neil G. Carey. Ninth edition. Alaska Northwest, 
Edmonds, Washington. 95 pp., illus. + map. U.S. $9. 95 
plus U.S.$2 postage; Cdn.$12.65. 



Biology and the mechanics of the wave-swept 
environment. 1988. By Mark W. Denny. Princeton 
University Press, Princeton, xiv + 329 pp., illus. Cloth 
U.S.$60; paper U.S.$25. 



Hawaii: the island of life. 1988. By Gavin Daws. Nature 
Conservancy of Hawaii (distributed by Pubhshers Group 
West, Emeryville, California). 156 pp., illus. U.S.$29.95. 

Islands in a far sea: nature and man in Hawaii. 1 988. By 
John L. Culliney. Sierra Club Books, San Francisco, xiv 
+ 410 pp., illus. U.S.$24.95. 

Living in a chemical world: occupational and 
environmental signiflcance of industrial carcinogens. 

1988. Edited by Cesare Maltoni and Irving J. Selikoff. 
New York Academy of Sciences, New York, xxv + 1045 
pp., illus. U.S.$260. 

tMountains: a natural history and hiking guide. 1989. By 
Margaret Fuller. Wiley, Somerset, New Jersey, xv + 255 
pp., illus. U.S.$12.95. 



Chambers biology dictionary. 1989. Edited by Peter 
M. B. Walker. Cambridge University Press, New York. 
c256 pp., illus. Cloth cU.S.$34.50; paper cU.S.$14.95. 

Effective risk communication: the role and responsibility 
of government and nongovernment organizations. 1 989. 
Edited by Vincent T. Covello, David B. McCallum, and 
Maria Pavlova. Plenum, New York. 365 pp. U.S. $85. 

The evolution of complexity by means of natural 
selection. 1988. By John Tyler Bonner. Princeton 
University Press, Princeton, xii + 260 pp., illus. Cloth 
U.S.$40; paper U.S.$13.95. 

The gene: a critical history. 1989. By Elof Axel Carlson. 
Iowa State University Press, Ames. 316 pp., illus. 
cU.S.$18.95. 



166 



The Canadian Field-Naturalist 



Vol. 104 



A history of biology to about the year 1900: a general 
introduction to the study of living things. 1989. By 
Charles Singer. Iowa State University Press, Ames. 616 
pp., iUus. cU.S.$22.95. 

*Lewis and Clark: pioneering naturaUsts. 1989. By Paul 
Russell Cutright. Reprint of 1969 edition. University of 
Nebraska Press, Lincoln, xvi + 506 pp., illus. U.S. $14.95. 

t Matrix population models: construction, analysis, and 
interpretation. 1989. By Hal Caswell. Sinauer, Sunder- 
land, Massachusetts, xiv + 328 pp., illus. Cloth U.S. $50; 
paper U.S.$28.95. 

My Serengeti years: the memoirs of an African game 
warden. 1988. By Myles Turner. Norton, New York, 
xxii + 221 pp., illus. U.S.$17.95. 

*No woman tenderfoot: Florence Merriam Bailey, pioneer 
naturalist. 1989. By Harriet Kofalk. Texas A and M 
University Press, College Station, xix + 225 pp., illus. 
U.S.$19.95. 

Books for Young Naturalists 

t Adventures in life sciences: process-oriented activities for 
grades 4-6. 1987. By Margy Kuntz. Fearon Teaching 
Aids. David S. Lake, Belmont, California. 43 pp., illus. 
U.S.$0.49. 

*Animal parenting. 1989. By Tony Seddon. Facts on 

File, New York. 62 pp., illus. U.S.$ 13.95. 

*The bug book and the bug bottle. 1987. By Hugh 
Danks. Somerville House Press, Toronto. 64 pp., illus. + 
bug bottle. $9.95. 

Do not disturb: the mysteries of animal hibernation and 
sleep. 1989. By Margery Facklam. Sierra Club Books/ 
Little, Brown, Boston. 47 pp., illus. U.S.$12.95. 



Experimenting with a microscope. 1988. By Maurice 
Bleifeld. Watts, New York. 110 pp., illus. U.S.$11.90. 

The garden book and the green house. 1989. By Wes 
Porter. Somerville House, Toronto. 64 pp., illus. + 
greenhouse. $10.95. 

The grandpa tree. 1988. By Mike Donahue. Roberts 
Rinehart, Boulder, Colorado. 22 pp., illus. U.S.$3.95. 

''Introducing birds to young naturalists. 1989. By Ilo 
Hiller. Texas A and M University Press, College Station. 
80 pp., illus. Cloth U.S.$21.50; paper U.S.$12.95. 

Keepers of the earth: native American stories and 
environmental activities for children. 1988. By Michael 
J. Caduto and Joseph Bruchac. Fulcrum, Golden, 
Colorado, xxv + 209 pp., illus. U.S.$ 18.95. 

Pond and river. 1988. By Steve Parker. Knopf, New 
York. 64 pp., illus. U.S.$12.95. 

The rock pool. 1988. By David Bellamy. Potter 
(Crown), New York. 24 pp., illus. U.S. $9.95. 

Why on earth? 1988. By the National Geographic 
Society, Washington. 96 pp., illus. U.S. $7.95. 

The wild inside: Sierra Club's guide to the great indoors. 

1988. By Linda Allison. Sierra Club, San Francisco. 144 
pp., illus. U.S.$7.95. 



* assigned for review 
tavailable for review 



TABLE OF CONTENTS (concluded) 



Book Reviews 

Zoology: The Collins Guide to the Birds of Britain and Europe with North Africa and the Middle 
East — The CoUins Guide to the Birds of South-East Asia — Birds of the Middle East and 
North Africa: A Companion Guide — An Annotated Bibliography of the Pike, Esox lucius 
(Osteichthyes: Salmoniformes) — Sharks — Suivi de la peche sportive dans les eaux de la 
region de Montreal — Guide to the Otoliths of the Bony Fishes of the Northeast Atlantic — 
Biology of the Land Crabs — Analyses in Behavioral Ecology: A Manual for Lab and Field 146 

Botany: The Flora of the Tobermory Islands, Bruce Pensinsula National Park — Lichens of the 
Ottawa Region (Second Edition) — A Second Checklist and Bibliography of the Lichens and 
Allied Fungi of British Columbia — Mosses, Lichens and Ferns of Northwest North America 
— Illustrated Guide to Some Hornworts, Liverworts, and Mosses of Eastern Canada — 
Mushrooms and Other Fungi of the Midcontinental United States — How to Identify 
Mushrooms to Genus VI: Modern Genera 151 

Environment: Stones of Silence: Journeys to the Himalaya — Fire in America: A Cultural History of 
Wildland and Rural Fire — Symbiosis: An Introduction to Biological Association — 
Population Ecology of Individuals — Wetlands of Canada 157 

Miscellaneous: The Outermost House 162 

New Titles 162 

Mailing date of the previous issue 103(4) : 26 September 1990 



THE CANADIAN FIELD NATURALIST Volume 104, Number 1 1990 



Articles 

Rare and Endangered Fishes and Marine Mammals of Canada: 
COSEWIC Fish and Marine Mammal Subcomittee Status Reports: VI 

R. R. Campbell 1 

Status of the Fourhorn Sculpin, Myoxocephalus quadricornis, in Canada 

J. Houston 7 

Status of the Spoonhead Sculpin, Cottus ricei, in Canada J. HOUSTON 14 

Status of the Bering Wolffish, Anarhichas orientalis, in Canada 

J. Houston and D. E. McAllister 20 

Status of the Blackline Prickleback, Acantholumpenus mackayi, in Canada 

J. Houston and D. E. McAllister 24 

Status of the Margined Madtom, Noturus insignis, in Canada 

Cheryl D. Goodchild 29 

Status of the Brook Silverside, Labidesthes sicculus, in Canada 

Cheryl D. Goodchild 36 

Status of the Banded Killifish, Fundulus diaphanus, in Canada J. Houston 45 

Status of the Least Darter, Etheostoma microperca, in Canada Ken W. Dalton 53 

Status of the River Darter, Percina schumardi, in Canada Ken W. Dalton 59 

Status of the Redbreast Sunfish, Lepomis auritus, in Canada J. HOUSTON 64 

Status of the Orangespotted Sunfish, Lepomis humilis, in Canada 

Douglas B. Noltie 69 

Status of the Bigmouth Buffalo, Ictiobus cyprinellus, in Canada 

Cheryl D. Goodchild 87 

Status of the Black Buffalo, Ictiobus niger, in Canada J. HOUSTON 98 

Status of the Golden Redhorse, Moxostoma erythrurum, in Canada 

Cheryl D. Goodchild 103 

Status of Dall's Porpoise, Phocoenoides dalli, in Canada Thomas A. JEFFERSON 1 12 

Status of Blainville's Beaked Whale, Mesoplodon densirostrus, in Canada 

J. Houston 117 

Status of Hubbs' Beaked Whale, Mesoplodon carihubbsi, in Canada J. Houston 121 

Status of Sowerby's Beaked Whale, Mesoplodon bidens, in Canada 

Jon Lien and Frances Barry 125 

Status of Stejneger's Beaked Whale, Mesoplodon stejnegeri, in Canada J. Houston 131 

Status of True's Beaked Whale, Mesoplodon mirus, in Canada J. Houston 135 

Status of the Ringed Seal, Phoca hispida, in Canada MICHAEL C. S. KiNGSLEY 138 



concluded on inside back cover 



ISSN 0008-3550 



The CANADIAN 
FIELD-NATURALIST 



Published by THE OTTAWA FIELD-NATURALISTS' CLUB, Ottawa, Canada 



r// 







Vi^ 







Volume 104, Number 2 



April-June 1990 



The Ottawa Field-Naturalists' Club 

FOUNDED IN 1879 

Patron 

His Excellency The Right Honourable Ramon John Hnatyshyn, P.C., C.C., C.M.M., Q.C., 

Governor General of Canada 

The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada's natural 
heritage; to encourage investigation and publish the results of research in all fields of natural history and to diffuse 
information on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, 
maintaining or restoring environments of high quality for living things. 



Honorary Members 



Edward L. Bousfield 
Irwin M. Brodo 
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President: 
Vice-Presidents: 



Claude E. Garton 
W. Earl Godfrey 
C. Stuart Houston 
Louise de K. Lawrence 
Thomas H. Manning 
Don E. McAllister 



Stewart D. MacDonald Hugh M. Raup 



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Verna Ross McGiffin 
Hue N. MacKenzie 
Eugene G. Munroe 
Robert W. Nero 



1990 Council 



Jeff Harrison 



Roy John 
Don Cuddy 

Recording Secretary: Elizabeth Fox 

Corresponding Secretary: Eileen Evans 

Treasurer: Mike Scromeda 



Ronald E. Bedford 
Barry Bendell 
Steve Blight 
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Francis R. Cook 
Don Davidson 
Enid Frankton 
Deirdre Furlong 



Loris S. Russell 
Douglas B. O. Savile 
Pauline Snure 
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Bill Gummer 
Paul Hamilton 
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Kenneth Strang 
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LOP IJO 
La Roi, Department 



of Botany, 



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The Canadian Field-Naturalist 

The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists' Club. Opinions and ideas 
expressed in this journal do not necessarily reflect those of The Ottawa Field-Naturalists' Club or any other agency. 

Editor: Francis R. Cook, R.R. 3, North Augusta, Ontario KOG IRO; (613) 996-1755 

Assistant to Editor: P. J. Narraway; Copy Editor: Joyce C. Cook 

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University of Alberta, Edmonton, Alberta T6G 2E9 
Associate Editors: Anthony J. Erskine 

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Back Numbers and Index 

Most back numbers of this journal and its predecessors, Transactions of The Ottawa Field-Naturalists' Cluh, 1879- 
1 886, and The Ottawa Naturalist. \ 887- 1919, and Transactions of The Ottawa Field- Naturalists ' Club and The Ottawa 
Naturalist — Index compiled by John M. Gillett, may be purchased from the Business Manager. 

Cover: An adult female tundra Peregrine Falcon(Fa/co/jeregr/>iMi;wAj^nai)about to settle over eggs at an eyrie in the 
Kecwatin District of the Northwest Territories. Photograph courtesy of Gordon S. Court. See article pages 
255 272. 



W. Bruce McGillivray 
William O. Pruitt, Jr. 
Stephen M. Smith 
Constantinus G. Van Zyll de Jong 



The Canadian Field-Naturalist 



Volume 104, Number 2 



April- June 1990 



Peregrene Falcons in the 1980s 



Dedication: Joseph Hickey 




It is indeed a pleasure to dedicate this issue to Dr. 
Joseph Hickey. His dedication to the Peregrine 
Falcon and its conservation is unrivaled. In the late 
1930s he organized a survey of the species 
throughout eastern North America so that the data 
could "be consulted by those who wish to study and 
evaluate subsequent population changes of this 
interesting species." I doubt if anyone reading the 
account in The Auk in the spring of 1942 realised 
how interesting and important these data would be. 

The Madison Peregrine Conference of 1965 is too 
well known as a landmark for the environmental 
movement to need further comment. The proceed- 
ings, edited by Joe alone, have become a classic. His 
subsequent follow-up, with Dan Anderson, placed 
eggshell thinning in the peregrine and many other 
species on a firm statistical basis. Joe gave the 
opening address to the Sacramento Peregrine 
meeting, held twenty years after that at Madison. 
The comparison of the size of these two meetings 
says much for the growth of interest in environmen- 
tal affairs. To mention just one statistic, the 
Sacramento meeting required four editors. 

In his 1942 Auk article, Joe quotes that the 
ancient Persians deemed it necessary that a falconer 
"be good-tempered, pleasant spoken and of a 
cheerful and cherry countenance." Although not a 
falconer himself that description certainly applies to 
Joe Hickey. 

David B. Peakall 



167 



Prospects for the Peregrine Falcon, Falco peregrinus, 
in the Nineties 

David B. Peakall 

Canadian Wildlife Service, National Wildlife Research Centre, Ottawa, Ontario Kl A 0H3 

Peakall, David B. 1990. Prospects for the Peregrine Falcon, Falco peregrinus, in the nineties. Canadian 
Field-Naturalist 104(2): 168-173. 

The first continent-wide survey of Peregrine Falcons {Falco peregrinus) was taken in 1970 and was followed by 
additional surveys in 1975, 1980 and 1985. The results of those of the 1970s were previously published in The Canadian 
Field-Naturalist and the present issue summarizes data obtained in the 1980s. These studies combined provide the 
basis for assessment of the species' prospects in the nineties and for some suggestions on the emphasis for studies in the 
next decade. Ecological differences among three North American geographic races have contributed to their present 
differential success. F. p. pealei occurs and breeds along British Columbian and Alaskan coasts. F. p. tundrius breeds 
in the arctic and winters in Latin America. F. p. anatum occupied a broad band from interior Alaska south through 
western Canada and a narrow range in the central and eastern portions of the country. Of the three, F. p. anatum has 
been affected the most, and was extirpated in eastern Canada and the United States. F. p. pealei and F. p. tundrius 
seem secure enough that the costly five-year surveys may no longer be justified for them. F. p. anatum is still at risk and 
its former range has been the focus of re-introduction programs. The introduction of captive-bred stocks has met with 
more success in the United States than in Canada. The observation that many falcons are only seen for one season is 
disturbing. Allelic frequency and DNA fingerprinting would aid in determining the relative effect of different stocks 
used in the release programs. Levels of organochlorines are still a concern. Release sites should be carefully chosen and 
the analysis of abandoned eggs continued. Finally, the studies of residues in Latin America should take a broad 
approach to identify specific problems of pesticide usage. 

Le premier inventaire pancontinental du Faucon pelerin a ete effectue en 1970 et suivi par d'autres inventaires en 1975, 
1980 et 1985. Les resultats des inventaires des annees 70 ont deja ete publics dans un numero du Canadian 
Field-Naturalist et le present numero resume les donnees obtenues dans les annees 80. Regroupees, ces etudes 
fournissent un fondement pour revaluation des perspectives de I'espece pour les annees 90 et de quelques suggestions 
des points principaux a etudier pour la prochaine decennie. Les differences ecologiques entre trois races geographiques 
nord-americaines ont contribue au succes present. Le F. p. pealei se retrouve et se reproduit le long des cotes de la 
Colombie-Britannique et de I'Alaska. Le F. p. tundrius se reproduit dans I'Arctique et hiverne en Amerique latine. Le 
F. p. anatum occupe un large couloir, partant du sud de I'Alaska et traversant I'ouest du Canada, ainsi qu'une region 
plus etroite au centre et a Test du pays. Parmi les trois races de faucons, le F. p. anatum a ete le plus touche et etait 
disparu dans Test du Canada et aux Etats-Unis. La survie du F. p. pealei et du F. p. tundrius semble assuree et les 
inventaires couteux de cinq ans pourraient ne plus etre justifies. Toutefois, le F. p. anatum est encore en danger et son 
ancien territoire a ete le sujet de programmes d'introduction. L'introduction de Faucons eleves en captivite a eu plus de 
succes aux Etats-Unis qu'au Canada. Les observations que plusieurs faucons ne sont reperes que pour une saison de 
reproduction sont troublantes. La frequence des genes allomorphes et les etudes de traces d'ADN sont necessaires pour 
determiner I'effet relatif des differentes sources d'approvisionnement utilisees dans les programmes de lachers. Les 
niveaux de contaminants organochlores demeurent une inquietude. Les zones de mise en liberte des faucons devraient 
etre choisies avec soin et I'analyse d'organochlores dans les oeufs abandonnes, poursuivie. Enfin, I'analyse des niveaux 
de contaminants organochlores en Amerique latine devrait utiliser une approche plus vaste pour identifier les 
problemes particuliers a cet endroit. 

Key Words: Peregrine Falcon, Falco peregrinus, surveys, ecology, breeding, re-introductions, organochlorines. 

This is the third issue of The Canadian exerting their greatest influence on the terminal 

Field- Naturalist to be devoted primarily to studies predators. Nevertheless, we should be careful not 

on the Peregrine Falcon (Falco peregrinus). I am to take the peregrine as the sole litmus paper of 

both honoured and intimidated to follow in the environmental quality. It is likely that the 

foot-stepsof Joe Hickey and Ian Newton in writing peregrine's choice of food {see Noble et al. this 

a preface to such an issue. Birds of prey have been issue) is sufficiently catholic to buffer its 

considered good indicators of environmental population against many changes that may occur 

health because they represent the terminal focus of in its food-chain. The cliff-nesting peregrine is also 

food chains and thus are sensitive to any important likely to be largely unaffected by habitat change 

change within the ecosystem. Certainly this has such as that caused by acid rain. Climate change 

been the case lor the organochlorine pesticides may affect the peregrine in the Prairie Provinces; 

which bio-accumulated through the food chain one might hypothesize that the Prairie Falcon 

168 



1990 



Peakall: Prospects for the Peregrine Falcon 



169 



{Falco mexicanus) is likely to increase at the 
expense of the peregrine. 

However, the question to be examined here is 
"What are the prospects for the peregrine in 
Canada in the 1990s? rather than "What does the 
peregrine tell us about the Canadian 
enviroiiment?" 

The major factor in the decline of the peregrine 
has been the widespread use of organochlorine 
pesticides" As Cade (1968) expressed it: "Down 
through the centuries, not all the falcon trappers, 
egg collectors, war ministries concerned for their 
messenger pigeons, or misguided gunmen have 
been able to effect a significant reduction in the 
numbers of breeding falcons. But the simple 
laboratory trick of adding a few chlorine molecules 
to a hydrocarbon and the massive apphcation of 
this unnatural class of chemicals to the environ- 
ment can do what none of these other grosser, 
seemingly more harmful agents could do." 

The ecological differences between the three 
sub-species that occur in Canada are so marked 
that it is necessary to consider each separately. 

Falco peregrinus pealei is essentially resident 
along the coasts of British Columbia and Alaska. 
In the Queen Charlotte Islands, a major nesting 
area of this race, significant decrease in the 
population occurred from 1970 to 1975 and it has 
subsequently recovered (Munro and van Drim- 
melen 1988). While it is impossible to assign a 
definite cause, it can be pointed out that a 
significant proportion of eggs analysed from the 
late 1960s and early 1970s had levels of DDE above 
the critical range, (Peakall et al., this issue) and the 
residue levels of a small sample of eggs collected in 
1986 are much lower, comfortably below critical 
levels. Kiff ( 1 988), in his overview of the changes in 
status of the peregrine in North America, shows 
the range of F. p. pealei as the one area in which no 
significant decline occurred. Although other 
pressures can be listed, it appears that the future of 
F. p. pealei is dependent on the populations of its 
seabird prey along the Pacific coast of North 
America. Detailed studies of the population of the 
two chief prey species, Cassin's Auklet 
{Ptychoramphus aeutica) and Ancient Murrelet 
{Synthliboramphus antiquum) are not available, 
but apart from local declines of Cassin's Auklet 
around Langara Island, the populations appear to 
be in good shape (A. J. Gaston, personal 
communication). 

In contrast, F. p. tundrius is highly migratory, 
breeding in the Arctic and wintering in Latin 
America. The population of F. p. tundrius declined 
markedly during the 1960s and the decline may 
have started as early as the 1950s. A decline of 65% 
between the late 1960s and 1985 was estimated for 
Alaska (Ambrose et al. 1988). Kiff (1988) 



calculated a population of 448 pairs in 1985 for the 
entire North American Arctic and Sub-arctic zone 
compared to an estimated historical population of 
5000-8000 pairs. Comparison of the 1970, 1975 and 
1980 surveys indicates that recovery started 
between 1975-1980. A further improvement was 
noted in most areas between 1980 and 1985. The 
levels of organochlorines in the eggs of F. p. 
tundrius have decreased and only a small 
proportion have levels above those considered 
critical (Peakall et al., this issue). 

The data presented in this issue on the residue 
levels of potential prey species of Peregrine 
Falcons collected in Latin America shows 
generally low levels from Surinam and Costa Rica 
and higher levels from Peru and Ecuador (Fyfe et 
al., this issue). It is difficult to obtain data on the 
usage of pesticides in Latin America. Maltby 
(1980) estimated that the total usage of DDT in 
Latin America was 16 000 000 kg in 1978 and 
forecast a 25% decrease in the usage of this 
insecticide over the next decade. Four countries — 
Mexico, Brazil, Argentina and Columbia — 
accounted for 90% of the 1978 total. To put these 
figures in perspective, the total amount of DDT 
used in New Brunswick over the period 1952-1965 
was 5 750 000 kg and maximum annual use was 
600 000 kg. and the area of New Brunswick is a 
fifteenth of that of Columbia and less than one 
percent of Brazil. 

F. p. anatum occupied a broad band from the 
interior of Alaska south through much of western 
Canada with rather narrower range across the 
central and eastern parts of the country. Its 
migrations are much more limited than those of F. 
p. tundrius. It was extirpated in the eastern half of 
Canada and in the eastern United States. The re- 
introduction program into eastern Canada has 
been detailed by Holroyd and Banasch elsewhere 
in this issue. Critical evaluation of the success of 
the re-introduction programs is essential. 

The starting point of any evaluation of any 
aspect of the biology of the peregrine is the 
Sacramento meeting of November 1985 (Cade et 
al. 1988). In his thought-provoking summary to 
this conference Ian Nisbet (1988a) posed four 
"irreverent" questions: (1) What caused the 
population crashes?; (2) What is known about 
population dynamics of peregrines?; (3) What is 
being learned about the biology of the peregrine 
from captive breeding programs?; (4) Where have 
all the captive-reared birds gone? 

These questions were also considered by the 
editors — Tom Cade, Jim Enderson, Carl 
Thelander and Clayton White — in the conclusion 
of the proceedings of the meeting. The first question 
they only considered briefly and since it is an issue 
that Ian Nisbet and I debated at Sacramento (Nisbet 



170 



The Canadian Field-Naturalist 



Vol. 104 



1988b; Risebrough and Peakall 1988), I think we 
can leave this fascinating question with lan's final 
words "the critical evidence — direct measures of 
exposure during the periods of decline — is lacking 
for nearly all areas." The more relevant questions 
are: "Do we know the levels of pollutants that are 
critical to the peregrine?" and "Are these levels still 
occurring in peregrines in the wild?" I would submit 
that we can answer the first question fairly well for 
most chemicals of concern on an individual basis. 
The data on which this statement is based are 
reviewed elsewhere (Peakall et al. this issue). Studies 
on raptors, however, of combinations of pollutants 
are few. The answer to the second question is that in 
some eggs the level of DDE is still uncomfortably 
close to that at which effects might be expected, 
especially in view of our lack of knowledge of the 
effect of combinations of chemicals. Also, we lack 
knowledge concerning the chronic — multi- 
generation — effects on raptors. Analysis of the 
Canadian sightings of released peregrines reveals the 
disconcerting fact that pairs only return for a short 
time, usually only one season (Holroyd and 
Banasch, this issue). Population dynamics data 
strongly suggests that adult mortality of the 
peregrine is normally low. It is possible that the 
levels of organochlorines increase sufficiently 
between the first and second years of breeding to 
cause problems. There are no good time series of 
eggs of individual females from eastern North 
America, but data from California indicate that 
equilibrium levels of organochlorine residues are 
reached in the second year (Kiff and Peakall, 
unpublished data). However, it is Ukely that the 
problems caused would be loss of eggs — due to 
breakage — or infertility rather than mortality of 
adults. 

The editors of the Sacramento proceedings take 
a more positive approach to the second question 
than Nisbet, although, like, all researchers, they 
acknowledge the need for additional studies. In 
this context 1 would like to draw attention to the 
paper in this issue by Court and co-workers, and 
additional material published by this group 
elsewhere (Court et al. 1988, 1989). Apart from 
their importance as a contribution to our 
knowledge of the breeding biology of the 
peregrine, they raise an important question about 
the value of intensive — as opposed to extensive — 
surveys as the index of population in the Arctic 
where the harsh and variable climate can cause 
major annual changes in productivity. An 
evaluation of the need for extensive surveys is 
made later in this paper. 

For the third question the editors again have a 
more positive attitude than Nisbet. The list of 
papers given by the editors is quite impressive. 
Personally, I have doubts on the value of studying 



many of the aspects of breeding biology that Nisbet 
lists in captive birds. On the other hand, I agree 
with Nisbet that it is important to identify the 
characteristics that lead to the survival of captive- 
bred birds in the ecological conditions into which 
we release them. 

The last, and most critical, question posed by 
Nisbet is "Where have all the captive-reared bird 
gone?" It is this question that the Editors consider 
at greatest length. It is possible to come up with 
almost any answer when producing survivorship 
tables. From a Canadian perspective the data 
presented in Nisbet's only table, which compared 
the number of birds released with the number of 
breeding pairs found in 1985, is the most critical. In 
Canada, 588 released birds resulted in seven 
known breeding pairs, while for the eastern United 
States the 51 1 birds released resulted in 25 known 
breeing pairs. The pattern of the release figures for 
Canada (Fyfe 1988) is very similar to that for the 
eastern United States (compiled by the editors. 
Cade et al. 1988: p. 860). Thus, on the basis of the 
raw data, the Canadian program is only a quarter 
as successful as that in the eastern United States. 

Reasons can be considered under two main, but 
not mutually exclusive, headings. The first is that 
the Canadian release program does produce less 
independent peregrines than the corresponding 
U.S. program and second that it only appears to be 
so. There are several possible reasons why the 
Canadian program is less successful. Perhaps the 
captive breeding techniques and mode of release 
are sufficiently different to lead to differential 
mortality, but this does not appear to be so. As far 
as release sites are concerned, the ratio of urban to 
rural is similar, although towers are not used in 
Canada. Limited residue level data (Gilroy and 
Barclay 1988; Peakall et al. this issue) do not 
indicate that organochlorine residues are higher in 
Canada than in the United States. The harsher 
Canadian winter may force longer migration 
patterns leading to increased mortality. The gene 
pool of the released birds is known to be different. 
This question is examined in more detail later. 

There are also several factors which could make 
it appear that the Canadian program is less 
successful. Certainly it is possible that peregrines 
are breeding in Canada in areas remote from 
humans. The density of observers is much lower 
and access to possible breeding sites more difficult 
in Canada. As Joe said, nearly half a century ago 
(in formal terms, Hickey 1942), commenting on the 
fact that 67% of the 408 nesting sites were located 
in the United States, "this preponderance may 
simply reflect the intensity of field work in the 
central and eastern parts of the continent, rather 
than the geographic indication of the species' 
center of abundance." Happily both Joe and one of 



1990 



Peakall: Prospects for the Peregrine Falcon 



171 



the other major contributors to this survey — 
Walter Spofford — are still with us. 

Nevertheless, the number of potential breeding 
sites in southern Canada may be less than that in 
the north-eastern United States; certainly the 
population in the Maritimes was never large. 
Emigration to the U.S. may be greater than 
immigration from the U.S. The data on banded 
birds are too limited to make a case one way or the 
other. 

The gene pool used for captive breeding that 
provided birds for the release programs are 
markedly different. Temple (1988) presented data 
on the contributions of various sub-species to the 
gene pool of captive-reared peregrines that have 
been released in the eastern United States over the 
period 1975-1985. F. p. tundrius accounted for 
c45% and an equal contribution of c 1 8% was made 
by F. p. anatum and the southern European F. p. 
brookei. Temple suggests the name ''Falco 
peregrinus cadeV for this novel regional "sub- 
species". The Canadian release program used pure 
F. p. anatum stock although there was an effort to 
use as broadly based a stock as possible. Fyfe 
(1988) refers to twelve peregrine nestlings being 
collected from the wild in 1970 from eyries as far 
apart as the Mackenzie Valley and Labrador. 
Subsequently the gene pool was augmented by 
additional birds, mainly from the northern boreal 
forests. If we followed the lead of Stan Temple, the 
name of this "sub-species" should be ''Falco 
peregrinus fyfei. " 

These "names" are, of course, used most inform- 
ally here; under the Internatioal Code of Zoological 
Nomenclature, formal scientific names cannot be 
applied to hybrids, nor are these stocks subspecies in 
a formal nomenclatural sense. F.p. anatum does not 
receive a new name on introduction. 

Morizot (1988) has published preliminary 
account of the biochemical variability in peregrine 
populations. These data have been used to make 
estimates of the proportions of various popula- 
tions in migrants trapped at Assateague Island, 
Virginia and Padre Island, Texas. The findings 
suggest that the percentage of North Slope 
Alaskan birds in Virginia and of Greenland birds 
in Texas are higher than is suggested by banding 
data. Although the author mentions samples from 
the captive breeding colonies at Ithaca, New York 
and Wainwright, Alberta these data do not appear 
in tabulated data nor are they discussed in the text. 
It would be fascinating to know allele frequencies 
for "cadei" and ''fyfei". Detailed DNA studies of 
released birds would also be informative, but 
regrettably these techniques were not available 
when the releases program began fifteen years ago. 
Even at this stage it would be useful to have 
information on the allele frequencies of birds 



released in Canada and United States. In view of 
the marked differences in the gene pool used it 
should certainly be possible to determine the 
amount of interaction between the birds released in 
the two countries. A comparison between the 
success of the two stocks of such different genetic 
composition — one from a single sub-species of a 
population that had passed through a relatively 
narrow neck and the other that was broadly based 
involving several sub-species — released in 
comparatively similar areas might well have 
considerable theoretical interest and practical 
importance. One would suppose that variability in 
'fyfei'' would be markedly lower than in "cadei". 
While such restriction are viewed with some alarm, 
on the grounds that decreased variability gives the 
species a decreased chance of survival if faced with 
new conditions, it should be pointed out that the 
Elephant Seal (Mirounga angustirostris) passed 
through a narrow neck and since then the 
population has increased greatly despite its very 
low genetic diversity. 

Before World War II there were only a few cases 
recorded of peregrines nesting in big cities, most 
notably the famous Sun-Life pair in Montreal. 
Thus, the urbanization of the peregrine in eastern 
North America represents a major change in its 
ecology. One unanswered and indeed unanswerable 
question is whether this phenomenon would have 
occurred naturally as clusters of high buildings 
became the norm. Essentially there were no 
peregrines in eastern North America during the 
1950s and 1960s, and in other areas of the world the 
numbers were depressed so that colonization of new 
habitat would be unlikely. As Cade and Bird make 
clear in their article it is not merely that urban sites 
were used as release sites as there are instances of 
rural released birds entering the city to breed, and 
peregrines have colonized several cities in which no 
releases were made. They hypothesize that captive- 
raised birds are less afraid of humans and have more 
knowledge of artificial structures. If this is correct it 
will be interesting to see what future generations of 
peregrines do, although it will be impossible to know 
with any certainty what has been the role of such 
experience. 

The saga of the colonization of cities by the 
peregrine will be followed with great interest. At the 
moment it is not possible to say what is the true 
breeding successs of these urban peregrines. There is 
great interest in the fate of the birds, many are fed 
artificially, and many fledglings are rescued. Most of 
the interest is positive, but the pigeon fanciers have 
strong antagonism to the peregrine. Only when the 
interest in the species dies down will we really be able 
to see how well the peregrine is adapting to that most 
unnatural of all environments, the core of the big 
city. 



172 



The Canadian Field-Naturalist 



Vol. 104 



Another problem that needs a decision is how 
much and what type of census work should be 
carried out on the peregrine. In 1970, on the heels of 
the discovery that the peregrine population in 
eastern North America was gone and that the 
Alaskan populations, like many others throughout 
the northern temperate zone, were greatly reduced, 
the first North American peregrine survey of 
breeding pairs was organized. There can be no 
doubt about the need for this survey, it was essential 
to document the status of the species. It confirmed 
the suspicion that the peregrine was indeed in poor 
shape over wide areas of the Arctic. The 1975 survey 
confirmed this although there were signs that the 
downward movement of the population had been 
arrested. The 1980 and 1985 surveys, both published 
in this issue of The Canadian Field-Naturalist, 
confirm the findings from elsewhere (documented at 
the Sacramento meeting, Cade et al. 1988) that the 
status of the species has improved considerably. As I 
write, the plans for the 1990 survey are being 
implemented. It is hard to come up with a definite 
figure for the cost of this survey but it is in the tens of 
thousands of dollars. Certainly the time has come to 
examine the policy of five-year surveys. The amount 
of time and effort that is available for conservation 
projects is limited and the question "Is this the best 
use of money and human resources?" must be asked. 

Court et al. in this issue puts forward the idea that 
annual intensive studies are more cost effective than 
extensive surveys. The weakness of extensive 
surveys is that the results are highly dependent on 
the weather conditions of that particular year. While 
annual intensive studies get round this point, the 
extrapolation to the entire region from a few data 
points causes a good deal of uncertainty in the data. 
Further, although Court and co-workers argue that 
the data can be collected on only three visits, the cost 
of an annual program to obtain this data may well 
equal the five-year broad survey. 

Another censusing technique is based on sighting 
and trapping data on migration. Looking at the data 
from Assateague Island, Virginia, we see that 63 
peregrines were sighted in 1970, 145 in 1975, 369 in 
1980 and 483 in 1985 compared to an annual figure 
of 475 for the period 1939-1947 (Ward et al. 1988). 
In this type of data there are problems of re-sighting 
and, conversely, of birds not sighted and, with the 
exception of trapping of banded birds orsightings of 
color bands or dye, we know nothing of the origin of 
these birds. Nevertheless, the annual sightings from 
Assateague certainly reflect the population changes 
known from the five-year surveys. 

The migration pattern of the peregrine is highly 
complex. Yates et al. (1988) have plotted the 
recoveries of Canadian banded peregrines. Most of 
those banded in the Northwest Territories migrate 
over the Caribbean to South America, whereas 



some birds from the Yukon Territories, and most of 
the birds from Keewatin and northern Quebec, 
migrate via the east coast of the United States. The 
remainder of the birds from these areas migrate due 
south to the Gulf of Mexico. The bulk of the 
recoveries of birds banded in Alaska occurred at the 
Gulf of Mexico, although there is one recovery from 
the east coast. 

Following fairly closely on the heels of the 
pubUcation of the Sacramento Conference, one has 
to justify another volume on the peregrine. First, 
this issue of The Canadian Field-Naturalist has 
provided a vehicle for the publication of the 3rd and 
4th major surveys of the peregrine. From a 
Canadian perspective it provides the data base 
available on the organochlorine residues in both the 
peregrine and its prey and a comparison to other 
raptors. The data on the residues in peregrine prey 
from Latin America provides the first published 
survey of this type. The details of the Canadian 
release program had not been previously published 
and the review, from both sides of the longest 
undefended border in the world, of the rise of the 
urban peregrine is an important contribution. Based 
on all this information let me offer the following 
conclusions and recommendations: 

1). The health of the F. p. pealei and F. p. 
tundrius populations in Canada, as we enter the 
1990s, is good. Further extensive surveys in the 
arctic such as those undertaken in 1970, 1975, 1980 
and 1985 do not appear to be justified unless they are 
part of a broader program undertaken to study the 
biology of the north. 

2). The status of F. p. anatum in eastern Canada 
remains a source of concern. So far the release 
program has been apparently less successful than its 
American counterpart. A survey of eastern Canada, 
south of the tree-line, would be useful to determine 
if, in fact, the population is larger than known at 
present. Studies on frequency of alleles and 
fingerprinting of the DNA of peregrines in eastern 
North America would be valuable to determine the 
proportion of birds coming from the U.S. and 
Canadian release programs. 

3). The levels of organochlorines in peregrine 
eggs are still uncomfortably close to critical levels 
even in eastern Canada where it can be assumed that 
the birds do not migrate to Latin America although 
their prey may do so. The finding that many falcons 
are only seen at breeding sites for one summer is 
disturbing. While there is no evidence that pesticides 
are involved, it is possible that there is enough of an 
increase between the first and second years to cause 
failure. Care should be taken over the selection of 
release sites, especially for the large releases (20 to 30 
young per year per area) proposed by Holroyd and 
Banasch. For each such area studies should be 
carried on the initial releases to determine the 



1990 



Peakall: Prospects for the Peregrine Falcon 



173 



principal prey selected and analysis for organochlo- 
rines should be carried out on these species. Only 
those areas, in which it is demonstrated that the 
levels of organochlorines in prey are below critical 
levels, should be used for mass releases. Most 
shorebirds in the Bay of Fundy have been shown to 
have low organochlorine residue levels, whereas 
residues levels are still high in gulls in the Great 
Lakes and along the St. Lawrence. No deliberate 
collection of peregrine eggs is proposed, but the 
analysis of levels in abandoned eggs in the east 
should be continued. Current evidence suggests that 
the levels determined in this manner are typical of 
the population. 

4). Additional studies on the levels of organoch- 
lorines in wildlife in Latin America would be of 
value because of the very Umited data now available. 
However, this should not be geared specifically 
towards peregrine prey, but rather take a broader 
approach with the idea of identifying specific 
problems in Latin America. 

Literature Cited. 

Ambros R. E., R. J. Ritchie, C. M. White, R. F. Schempf, 

T. Swem and R. Dittrick. 1988. Changes in status of 
Peregrine Falcon populations in Alaska. Pages 73-82 in 
Peregrine Falcon populations: their management and 
recovery. Edited by T. J. Cade, J. H. Enderson, C. G. 
Thelander and C. M. White. The Peregrine Fund, Inc., 
Boise, Idaho. 

Cade, T. J. 1968. The Gyrfalcon and falconry. Living 
Bird 7: 237-240. 

Cade, T. J., J. H. Enderson, C. G. Thelander, and C. M. 
White. 1988. Peregrine Falcon populations: their 
management and recovery. The Peregrine Fund, Inc., 
Boise, Idaho. 949 pages. 

Court, G. S., C. C. Gates, and D. A.Boag. 1988. Natural 
history of the Peregrine Falcon in the Keewatin District 
of the Northwest Territories. Arctic 41: 17-30. 

Court, G. S., D. M. Bradley, C. C. Gates, and D. A. Boag. 
1989. Turnover and recruitment in a tundra Peregrine 
Falcon Falco peregrinus population. Ibis 131: 487-496. 

Fyfe, R. W. 1988. The Canadian Peregrine Falcon 
recovery program, 1967-1985. Pages 1121-11% in 
Peregrine Falcon populations: their management and 
recovery. Edited by T. J. Cade, J. H. Enderson, C. G. 
Thelander and C. M. White. The Peregrine Fund, Inc., 
Boise, Idaho. 

Gilroy, M. J., and J. H. Barclay. 1988. DDE Residues 
and eggshell characteristics of reestablished peregrines 
in the eastern United States. Pages 403-412 /« Peregrine 
Falcon populations: their management and recovery. 
Edited by T. J. Cade, J. H. Enderson, C. G. Thelander 
and C. M. White. The Peregrine Fund, Inc., Boise, 
Idaho. 

Hickey, J. J. 1942. Eastern population of the duck hawk. 
Auk 59: 176-204. 



Kiff, L. F. 1988. Commentary — Changes in the status of 
the Peregrine in North America: an overview. Pages 
123-140 in Peregrine Falcon populations: their 
management and recovery. Edited by T. J. Cade, J. H. 
Enderson, C. G. Thelander and C. M. White. The 
Peregrine Fund, Inc., Boise, Idaho. 

Maltby, C. 1980. Report on the use of pesticides in Latin 
America. UNIDO/IOD.353. 139 pages. 

Morizot, D. C. 1988. Biochemical genetic variability in 
Peregrine Falcon populations. Pages 773-778 in 
Peregrine Falcon populations: their management and 
recovery. Edited by T. J. Cade, J. H. Enderson, C. G. 
Thelander and C. M. White. The Peregrine Fund, Inc., 
Boise, Idaho. 

Munro, W. T., and B. van Drimmelen. 1988. Status of 
peregrines in the Queen Charlotte Islands, British 
Columbia. Pages 69-72 in Peregrine Falcon popula- 
tions: their management and recovery. Edited by T. J. 
Cade, J. H. Enderson, C. G. Thelander and C. M. 
White. The Peregrine Fund, Inc., Boise, Idaho. 

Nisbet, I. C. T. 1988a. Summary. Pages 851-856 In 
Peregrine Falcon populations: their management and 
recovery. Edited by T. J. Cade, J. H. Enderson, C. G. 
Thelander and C. M. White. The Peregrine Fund, Inc., 
Boise, Idaho. 

Nisbet, \. C. T. 1988b. The relative importance of DDE 
and dieldrin in the decline of Peregrine Falcon 
populations. Pages 351-376 in Peregrine Falcon 
populations: their management and recovery. Edited by 
T. J. Cade, J. H. Enderson, C. G. Thelander and C. M. 
White. The Peregrine Fund, Inc., Boise, Idaho. 

Risebrough, R. W., and D. B. Peakall. 1988. Commen- 
tary — The relative importance of the several 
Ooganochlorine in the decline of Peregrine Falcon 
populations. Pages 449 462 in Peregrine Falcon 
populations: their management and recovery. Edited by 
T. J. Cade, J. H. Enderson, C. G. Thelander and C. M. 
White. The Peregrine Fund, Inc., Boise, Idaho. 

Temple, S. A. 1988. Future goals and needs for the 
management and conservation of the Peregrine Falcon. 
Pages 843-848 in Peregrine Falcon populations: their 
management and recovery. Edited by T. J. Cade, J. H. 
Enderson, C. G. Thelander and C. M. White. The 
Peregrine Fund, Inc., Boise, Idaho. 

Ward, P. F., K. Timus, W. S. Seegar, M. A. Yates, and 
M. R. Fuller. 1988. Autumn migrations of Peregrine 
Falcons at Assateague Island, Maryland/ Virginia, 
1970-1984. In Peregrine Falcon populations: their 
management and recovery. Edited by T. J. Cade, J. H. 
Enderson, C. G. Thelander and C. M. White. The 
Peregrine Fund, Inc., Boise, Idaho. 

Yates, M. A., K. E. Riddle, and F. P. Ward. 1988. 
Recoveries of Peregrine Falcons migrating through the 
Eastern and Central United States, 1955-1985. Pages 
471-484 in Peregrine Falcon populations: their 
management and recovery. Edited by T. J. Cade, J. H. 
Enderson, C. G. Thelander and C. M. White. The 
Peregrine Fund, Inc., Boise, Idaho. 

Received 12 November 1989 
Accepted 28 January 1990 



The 1980 North American Peregrine Falcon, 
Falco peregrinus, Survey 

Clayton M. White', Richard W. Fyfe2, and David B. Lemon^ 

'Zoology Department, Brigham Young University, Prove, Utah 84602. 

2P.0. Box 3263, Ft. Saskatchewan, Alberta T8L 2T2. 

^General Delivery, Topsail, Conception Bay South, Newfoundland AOA 3Y0. 

White, Clayton M., Richard W. Fyfe, and David B. Lemon. 1990. The 1980 North American Peregrine Falcon, 
Falco peregrinus, survey. Canadian Field Naturalist 104(2): 174-181. 

Most areas of North America were surveyed for nesting of the Peregrine Falcon {Falco peregrinus) in 1980. Specific 
areas not adequately surveyed include the Aleutian Islands, southeast Alaska, Greenland, Labrador and Mexico. 
Compared to the 1975 North American surveys, many regional populations in 1980 had higher numbers of occupied 
sites and higher site productivity. There appeared to be a complete population recovery of Peregrine Falcons along 
portions of the Yukon River. Areas adjacent to the Yukon River, however, have remained stable at low levels or have 
declined further compared to previous survey results. The Mackenzie River Valley population appears to have 
remained stable whereas the population on the Mackenzie River delta has continued to decline. The arctic populations 
showed a trend toward stability in some areas whereas other areas had slight increases or declines since 1975. The 
Pacific maritime populations have remained stable since 1975 and population levels there are either near capacity or 
have already recovered to former numbers. No Peregrine Falcons were observed in southern Alberta, Saskatchewan, 
Manitoba, Ontario or Nova Scotia. Southern Quebec and northern Alberta had a total of 1 1 pairs of nesting Peregrine 
Falcons. The eastern United States had three nesting pairs in 1980 compared to no known pairs in 1975. The western 
United States and Mexico had at least 185 occupied sites compared to 62 in 1975. The increase observed in the western 
United States and Mexico is primarily a result of more search effort. The North American Peregrine Falcon 
population has stabilized since the population decline was identified in the 1960s. However, population increases or 
declines are continuing on a local level. 

En 1980, on a etudie la nidification du Faucon pelerin {Falco peregrinus) dans la plupart des regions de I'Amerique du 
Nord. Certaines regions n'ont pas ete etudiees suffisamment; ce sont les Ties Aleoutiennes, le sud-est de I'Alaska, le 
Greenland, le Labrador et le Mexique. Comparativement aux enquetes nord-americaines de 1975, les donnees 
demontrent qu'en 1980, plusieurs populations regionales occupaient un plus grand nombre de sites et avaient un taux 
de reproduction plus eleve. II semble y avoir eu un retablissement complet de la population de Faucons pelerins le long 
de certaines parties du fleuve Yukon. Les donnees des regions adjacentes au fleuve Yukon sont toutefois demeurees 
stables, a des niveaux bas, ou ont encore decline en comparaison avec les resultats du releve precedent. La population 
de la vallee du fleuve Mackenzie semble etre demeuree stable tandis que la population du delta du fleuve Mackenzie a 
continue de decliner. Les populations arctiques demontrent une tendance a la stabilite dans certaines regions tandis 
que dans d'autres, elles se sont accrues ou elles ont diminue depuis 1975. Les populations de la cote du Pacifique sont 
demeurees stables de puis 1975 et les niveaux des populations a cet endroit sont presque retablis ou ils ont deja atteint 
les nombres precedents. Aucun Faucon pelerin n'a ete observe au sud de I'Alberta, en Saskatchewan, au Manitoba, en 
Ontario ou en Nouvelle-Ecosse. Au sud du Quebec et au nord de I'Alberta, on a releve un total de 1 1 couples de 
Faucons pelerins nicheurs. L'est des Etats-Unis comptait trois couples nicheurs en 1980 tandis qu'en 1975 on n'en avait 
releve aucun. L'ouest des Etats-Unis et le Mexique ont au moins 185 sites habites comparativement a 62 en 1975. 
L'augmentation observee dans l'ouest des Etats-Unis resuhe principalement de recherches plus poussees. La 
population nord-americaine de Faucons pelerins s'est stabilisee depuis que son declin a ete reconnu au cours des 
annees 60. Des augmentations ou des diminutions de populations continuent toutefois a I'echelle regionale. 

Key Words: Peregrine Falcon, Falco peregrinus. Survey, North America. 

The MadLson Peregrine Conference in 1965 first (Fyfe et al. 1976) surveys indicated that migratory 

documented the severity of the Peregrine Falcon peregrine populations continued to decline 

(Falco peregrinus) population decline in North throughout most of their northern and interior 

America (Hickey 1969). This resulted in an effort ranges. The relatively sedentary population of 

to determine the extent of the population decline. Pacific marine peregrines remained stable. 

In 1969, a raptor conference held at Cornell The surveys reported herein represent a 

University reviewed the most recent field data on considerable increase in effort to determine the 

peregrine populations and recommended that a abundance, distribution and reproductive success 

North American survey of peregrine eyries be of the North American peregrine population 

conducted every five years beginning in 1970. The compared to the 1970 and 1975 surveys. The 

results of the 1970 (Cade and Fyfe 1970) and 1975 increased effort reflects an overall public 

174 



1990 



White, Fyfe, and Lemon: 1980 Peregrine Falcon Survey 



175 



awareness and support of peregrine related 
projects. This has resulted in peregrine conserva- 
tion and management projects receiving funding 
from government and non-government agencies, 
and the support of volunteers. 

This paper is a brief review of the 1980 surveys 
and will provide wildlife researchers and managers 
with essential information on the North American 
peregrine population in 1980. This survey is 
intended to complement the surveys reported in 
Cade and Fyfe (1970) and Fyfe et al. (1976). Cade 
et al. (1988) provide a thorough review of North 
American Peregrine Falcon populations, includ- 
ing results from this study and more recent data. 

Methods 

The 1980 survey in Canada was coordinated by 
the Canadian Wildlife Service who provided 
guidelines for collecting and recording data. The 
same guidelines were provided for United States 
participants. The peregrine surveys were con- 
ducted using a variety of methods including 
automobile, boat, helicopter and fixed-wing 
aircraft. Ground checks of nest sites were made 
when conditions allowed. Nest sites were ground 
checked using noise makers to cause the birds to fly 
from the site, climbing to sites or observing sites 
with binoculars or spotting scopes. 

In this paper, a nest site is defined as any location 
where peregrines have been observed and a nesting 
attempt has been documented. An occupied site is 
defined as a site occupied by a pair of adults or a 
single adult. 

Non-essential data from individual surveys has 
been omitted to keep the focus of the paper on the 
North American peregrine population. However, 
the names and addresses of the contributors for 
each survey are provided in the acknowledgments 
for reference purposes. 

Due to the length of time between the 1980 
surveys and the publication of these results, 
outdated material has been omitted and previously 
published material has been either omitted or cited 
appropriately. The original manuscript (80 pages 
of text plus 40 tables) containing detailed 
information is on file with the first two authors and 
the U.S. Fish and Wildlife Service, Office of 
Endangered Species, Washington, D.C. 20240. 
Specific data can be obtained from that 
manuscript. The literature cited in that manuscript 
was comprehensive. 

Results and Discussion 

The results of the peregrine surveys conducted in 
1 980 are presented in Table 1 . The North American 
overview of the 1980 survey results that follows is 
based on five regions; the Arctic tundra, the boreal 
forest north of 60° N latitude, the boreal forest 



south of 60°N latitude, the Pacific northwest, and 
the continental United States (excluding Alaska) 
and Mexico. 

For a thorough overview of the peregrine status 
in North America, which comprises the survey 
efforts conducted prior to 1980, the 1980 surveys 
reported in this paper, and more recent survey 
results, the reader should refer to Kiff (1988). 

The Arctic Tundra Region 

This region comprises the Canadian Arctic, 
Greenland and the arctic portions of Alaska. 

The number of occupied peregrine sites in the 
Canadian Arctic has recently increased as a result 
of numerous raptor surveys associated with 
environmental impact assessment projects in 1976 
and 1977. These recent surveys located approxi- 
mately 100 new sites which double the total 
number of known sites for the interior barrens. 
Calef and Heard (1979) presented results of two 
surveys that indicated high reproduction and site 
occupancy rates in the Canadian Arctic. In 1980, 
concerted efforts were made to survey the recently 
located sites, in particular the new sites in the 
Wager Bay area. As a result, the 1980 surveys of the 
arctic barrens and coastal areas were more through 
compared to previous surveys in these areas. 
Therefore the surveys presented in this paper are 
considered to be more representative of the arctic 
peregrine population than estimates determined 
from previous surveys. 

Most of the 1980 surveys in the Canadian Arctic 
were adversely affected by poor weather condi- 
tions throughout the summer following a later 
than usual arrival of spring. There were many 
surveys delayed or aborted due to weather. 
However, survey crews were confident of the 
reliabihty of the results they managed to collect 
and only presented those results. For example, 
during the survey of the Wager Bay area, 30 of the 
37 known sites were visited but at only 21 of the 
sites were the survey results satisfactory. 
Consequently, only 21 sites are included in the 
survey. 

The peregrine populations on the central coast, 
interior barrens and Ungava have remained 
relatively stable with 40-45% site occupancy. The 
Thelon River results indicate a slight population 
increase from previous surveys. 

Local peregrine population declines in the arctic 
have not entirely stopped. On the Yukon North 
Slope west of the Mackenzie River delta only two 
single adult peregrines were observed at 16 known 
sites. On Baffin Island, peregrines were not present 
at any of six sites. The 1980 Wager Bay survey also 
indicated a population decHne whereas earlier 
studies in the same area indicated the population 
was stable with high nest productivity (Calef and 
Heard 1979). The 1980 Wager Bay survey 



176 



The Canadian Field-Naturalist 



Vol. 104 



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The Canadian Field-Naturalist 



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indicated a decrease in site occupancy from 
approximately 59% to 38% since the 1976-77 
surveys conducted by Calef and Heard (1979). 

The west Greenland peregrine population 
appears to have remained stable since 1972 with an 
annual site occupancy rate of approximately 60%. 
The number of known sites in west Greenland has 
increased by approximately 60% since 1972. 

In the Alaskan Arctic, only 29% of the historical 
sites surveyed on the Colville River were 
productive but when new sites in addition to 
historical sites are considered, approximately 57% 
of all sites produced young. Other areas in the 
Arctic had similar site occupancy rates and survey 
results suggest the peregrine population has 
stabilized. 

More recent peregrine population estimates and 
population trends for this region can be found in 
Mattox and Seegar (1988) for Greenland; Falk and 
Moller (1988) for south Greenland; Bird and 
Weaver (1988) for Ungava Bay, Quebec; Bromley 
(1988) for Kitikmeot, Baffin and Keewatin, 
Northwest Territories; Mossop (1988) for the 
Yukon; Ambrose et al. (1988) for Alaska. 

The Boreal Forest Region North of60°N 

This region comprises areas of central and 
northern Alaska, the Yukon and Northwest 
Territories below treeline and north of 60° N 
latitude. 

The results of the 1980 peregrine surveys in this 
region indicated the beginning of a population 
recovery when compared to previous surveys. The 
most extensive recovery was documented along the 
Yukon River. The Yukon Territory populations 
recovered dramatically from apparently low levels 
in 1977. This recovery was aided by an 
immigration of peregrines moving up the Yukon 
River from Alaska as determined by band 
recoveries. Forty-eight peregrine nest sites have 
been located along the lower Yukon River in 
Alaska since 1977. This represented approximately 
30% of the total known sites in Alaska. The 
peregrine population along the lower Yukon 
River, Alaska, was very productive and accounted 
for 36% of the young produced in Alaska in 1980. 
In contrast, the Tanana River peregrine popula- 
tion in Alaska had only a 20% site occupancy rate. 
A new Peregrine nest site was located near the 
effluent of the Copper River, an area where 
peregrines were not previously known to nest. 

Peregrine populations along the Porcupine and 
Mackenzie Rivers either showed an increase in site 
occupancy rate or remained stable. Recent data 
from the Peel River and Campbell Hills 
(Mackenzie River delta) indicated a severe 
population decline possibility as a result of 
reproductive failure and decreased site occupancy 
rate. 



More recent peregrine population estimates and 
population trends for this region can be found in 
Bromley and Matthews (1988) for the Mackenzie 
River Valley, Northwest Territories; Mossop 
(1988) for the Yukon; Ambrose et al. (1988) for 
Alaska. 

The Boreal Forest Region South of60°N 

This region comprises areas of mainland 
Canada south of 60° N latitude excluding Pacific 
coastal areas. 

The 1980 surveys conducted in the boreal forest 
regions south of 60° N were the most complete to 
date, although historical sites in Labrador and 
New Brunswick were not surveyed. Cade and Fyfe 
(1970) and Fyfe et al. (1976) reported a near 
complete disappearance of peregrines for this 
region in 1970 and 1975. The 1980 surveys 
investigated 141 of the 150 known sites in this 
region in addition to many areas of suitable habitat 
in Quebec, Saskatchewan and Manitoba. No 
peregrines were observed in southern Alberta, 
Saskatchewan, Ontario, Manitoba or Nova 
Scotia. 

Eleven sites were active in 1980 in northern 
Alberta and southern Quebec. Another historical 
site in southern Quebec was occupied in 1979. 
Eight of the peregrines observed at these sites in 
1979 and 1980 were raised in captivity and released 
from the Cornell University and Camp Wainw- 
right captive breeding facilities. Many of the other 
peregrines occupying these sites were also banded 
but their origin could not be determined. One 
peregrine occupying a site was released in 1977 in 
Edmonton, Alberta. 

The Pacific Northwest Maritime Region 

This region comprises southeast Alaska, the 
Aleutian Islands and the Queen Charlotte and Gulf 
Islands off the coast of British Columbia. 

Although the surveys conducted in the Aleutian 
Islands area were incomplete, the results suggested 
that the number of nesting peregrines there is large 
and productivity appears healthy. At least 33 nest 
sites were located in southeast Alaska between 
1978 and 1981 on coastal islands and on the 
mainland. In addition, 36 coastal cliffs on outer 
islands were occupied. Eight sites in southeast 
Alaska were located in tree cavities and one 
appeared to be in an Accipiter sp. nest. The 
locations of the tree cavity nesting peregrines 
indicates a 300 400 km northward extension of 
tree nesting peregrines that were reported in British 
Columbia by Campbell et al. (1977). There is a 
possibility that a large population of tree nesting 
peregrines exists in southeast Alaska. 

The peregrine population on the coast of British 
Columbia appeared to remain stable. The lack of 
obvious population fluctuations in this area 
indicated that the nesting density of peregrines 



1990 



White, Fyfe, and Lemon: 1980 Peregrine Falcon Survey 



179 



may be near its maximum. Survey results from 
Langara Island have remained consistent. The 
remaining areas of the Queen Charlotte Islands 
surveyed indicated that a definite population 
increase is occurring. The site occupancy in the 
Queen Charlotte Islands was approximately 67% 
and in the Gulf Islands, approximately 56%. 

More recent peregrine population estimates and 
population trends for this region can be found in 
Munro and van Drimmelen (1988) for British 
Columbia; Ambrose et al. (1988) for Alaska. 

The United States (excluding Alaska) 
and Mexico Region 

This region comprises the continental United 
States (excluding Alaska) and northwest portions 
of Mexico. 

The peregrine population in the continental 
United States east of the Mississippi River 
consisted of three nest sites in 1980. Two of the 
three pairs consisted of captive bred and released 
birds nesting on made-made towers in New Jersey 
and the third pair was of undetermined origin in 
coastal Maine. 

The peregrine population in the continental 
United States west of the Mississippi River 
consisted of 145 pairs and at least 185 occupied 
sites. This compares to 62 pairs in 1975. The 
apparent increase in 1980 was thought to reflect an 
increase in survey effort rather than being 
indicative of a population recovery. Many of the 
newly discovered occupied sites were located in 
California, Arizona, New Mexico and Mexico. 
The 1 1 new occupied sites located in California and 
the greater number of occupied sites in Mexico 
were clearly the results of increased survey effort. 
The increase in numbers in Baja and the Gulf of 
California area may have represented a true 
population recovery in addition to increased 
survey efforts. The peregrine populations in 
Colorado continued to show signs of decline even 
though two new occupied sites were located. 

More recent peregrine population estimates and 
population trends for this region can be found in 
Enderson et al. (1988) for Colorado; Ellis (1988) 
for Arizona; Walton et al. (1988) for California, 
Oregon, Washington and Nevada; Porter et al. 
(1988) for Baja California and the Gulf of 
California, Mexico; Hunt et al. (1988) for Texas 
and Northern Mexico. 

Acknowledgments 

The list of principal contributors to the 1980 
surveys is a long and impressive one which reflects 
the truely North American focus of the surveys. 
However, due to the number of years that have 
elapsed between 1980 and the publication of these 
results, much of the information concerning 
current addresses of the contributors is out of date. 



The address of the first contributor of each survey 
is provided for correspondence purposes. 

Principal Contributors of Survey Results: 

West Greenland: W. G. Mattox, Ohio Depart- 
ment of Natural Resources, Columbus, Ohio 
43229; W. A. Burnham, F. P. Ward, W. S. Seegar. 
Ungava Bay, Quebec/NWT: D. M. Bird, Macdo- 
nald Raptor Research Centre, Ste. Anne de 
Bellevue, Quebec H9X ICO; J. D. Weaver. South 
Baffin Island, NWT: G. Court and P. Trefry, 
Canadian Wildlife Service, Edmonton, Alberta 
T6B 2X3. Rankin Inlet, NWT: G. L. Erickson, 
Alberta Fish and Wildlife Division, Edmonton, 
Alberta T6H 4P2; C. Gates. Wager Bay, NWT: 
G. L. Erickson, Alberta Fish and Wildlife 
Division, Edmonton, Alberta T6H 4P2; D. 
Jacobs. Thelon River and Interior Barrens, NWT: 
L. Dickson and V. Stringer, Canadian Wildlife 
Service, Edmonton, Alberta T6B 2X3; G. 
Erickson, R. BulUon, J. and J. Faulkner, R. 
Moore, R. Turner. Anderson and Horton Rivers, 
NWT: T. Barry, Canadian Wildlife Service, 
Edmonton, Alberta T6B 2X3. Central Arctic 
Coast, NWT: H. Armbruster, Canadian Wildlife 
Service, Edmonton, Alberta T6B 2X3; K. Lloyd. 
Victoria and Banks Islands, NWT: G. AUiston, 
LGL Ltd., Toronto, Ontario M4R lAl; L. 
Dickson, U. Banasch. Yukon North Coast: D. 
Mossop and R. Hayes, Yukon Wildlife Branch, 
Whitehorse, Yukon Y 1 A 2C6. Central and Eastern 
Arctic Slope, Alaska: D. Roseneau, P. Bente and 
A. Springer, LGL Alaska Research Associates, 
Fairbanks, Alaska 99708. Colville River, Alaska: 
R. Ambrose, U.S. Fish and Wildlife Service, 
Anchorage, Alaska 99503. Mackenzie River 
Valley, NWT: L. Dickson, U. Banasch and G. 
Court, Canadian Wildlife Service, Edmonton, 
Alberta T6B 2X3. Porcupine River, Peel River, 
Yukon River and Alsek River, Yukon: D. Mossop 
and R. Hayes, Yukon Wildlife Branch, White- 
horse, Yukon YIA 2C6. Upper Yukon River, 
Alaska: R. Ambrose, U.S. Fish and Wildlife 
Service, Anchorage, Alaska 99503. Charley River, 
Alaska: R. Ambrose, U.S. Fish and Wildlife 
Service, Anchorage, Alaska 99503. Porcupine 
River, Alaska: R. Ritchie and J. Curatolo, Alaska 
Biological Research, Fairbanks, Alaska 99708. 
Middle Yukon River and Lower Yukon River, 
Alaska: A. Springer, P. Bente and D. Roseneau, 
LGL Alaska Research Associates Ltd., Fairbanks, 
Alaska 99708. Southwest Alaska: P. Mindell, 
Harvard University, Cambridge, Massachusetts 
02138; R. Dotson. Tanana River, Alaska: D. 
Roseneau, P. Bente and A. Springer, LGL Alaska 
Research Associates Ltd., Fairbanks, Alaska 
99708. Saskatchewan: S. Barber, M. Hlady and J. 
Kinnear, Tourism and Renewable Resources, 
Regina, Saskatchewan S4S 5W6; R. Rafuse. Nova 



180 



The Canadian Field-Naturalist 



Vol. 104 



Scotia: A. Smith, Canadian Wildlife Service, 
Sackville, New Brunswick; P. Barkhouse. 
Southern Quebec: D. Bird, Macdonald Raptor 
Research Centre, Ste. Anne de Bellevue, Quebec 
H9X ICO; M. Bureau, N. David, M. Lepage. 
Ontario: S. Bradley, 95 Arlington Avenue, 
Toronto, Ontario M6G 3C2. Lower Churchill 
River, Manitoba: W. Koonz, Department of 
Natural Resources, Winnipeg, Manitoba 
R3H 0W9; W. Anderson (deceased). Alberta 
north of 58° N: L. Johnston-Beaver, General 
Delivery, Roseneath, Ontario KOK 2X0. Alberta 
south of 58°N: Alberta Fish and Wildlife Division, 
Calgary, Alberta T2H OGl, O. Pall (deceased). 
Eastern United States: P. Nickerson, U.S. Fish and 
Wildlife Service, Newton Corner, Massachusetts 
02158. Upper Misissippi River, Lake Superior 
(north shore) and Border Lakes: P. Redig, College 
of Veterinary Medicine, University of Minnesota, 
St. Paul, Minnesota 55108. Washington: C. 
Anderson, Kirkland, Washington 98033; F. 
Dobler, S. Herman. Oregon: C. Henney, U.S. Fish 
and Wildlife Service, Corvallis, Oregon 97333. 
Montana: J. Summer, Arlee, Montana 59821. 
Idaho: R. Howard, U.S. Fish and Wildlife Service, 
Boise, Idaho 83705; M. Nelson. Wyoming: R. 
Oakleaf, Wyoming Game and Fish Department, 
Lander, Wyoming; M. Jenkins. Utah: A. Heggen 
and P. Wagner, Utah Division of Wildlife 
Resources, Salt Lake City, Utah 84116. Nevada: 
G. Herron, Nevada Department of Wildlife, Reno, 
Nevada 89520. Colorado: G. Craig, Colorado 
Division of Wildlife, Fort Collins, Colorado 
80526; J. Enderson. Arizona: D. Ellis, Patuxent 
Wildlife Research Centre, Laural, Maryland 
20708: J. Fackler. New Mexico: J. Hubbard, New 
Mexico Department of Game and Fish, Santa Fe, 
New Mexico 87503. California: D. Harlow, U.S. 
Fish and Wildlife Service, Sacramento, California; 
B. J. Wakon, C. G. Thelander, D. Boyce, G. 
Monk, M. Kirven, T. Scott, L. Kiff. Texas and 
northeast New Mexico: W. Hunt, Chihuahuan 
Desert Research Institute, Alpine, Texas 79830. 
Baja and Gulf of California, Mexico: R. Porter 
and M. Jenkins, U.S. Fish and Wildlife Service, 
Denver Federal Center, Denver, Colorado 80225; 
M. Kirven. Aleutian Islands-Alaska Peninsula: B. 
Day, Institute of Marine Sciences, University of 
Alaska, Fairbanks, Alaska 99775; P. Heglund, B. 
Reiwing, T. Early, E. Baily, D. Nysewander. 
Coastal and Inland Regions, Southeast Alaska: D. 
Van Horn, Malheur National Forest, John Day, 
Oregon 97845. Outer Coast, southeast Alaska: P. 
Schemph, U.S. Fish and Wildlife Service, Juneau, 
Alaska 99802. Vancouver and Gulf Islands, British 
Columbia: R. Davis, C. Elliot and G. Smith, B.C. 
Fish and Wildlife Branch, Nanaimo, British 
Columbia. Langara Island, British Columbia: 



R. W. Nelson, Camrose, Alberta T4V 2W6. Queen 
Charlotte Islands, British Columbia: K. Hodson 
and B. van Drimmelen, B.C. Fish and Wildlife 
Branch, Smithers, British Columbia VOJ 2N0. 

The following people are acknowledged for their 
contributions to the surveys: S. Ambrose, C. 
Anderson, D. Anderson, R. Banks, J. Bean, S. 
Belardo, D. Benfield, E. Boeker, W. Brewster, T. 
Conner, J. Dahlke, J. Egbert, L. Egbert, M. 
Fairchild. G. Flaxa, D. Forsell, M. Foster, F. 
Fridrikson, J. Fryxell, M. Fuller, D. Gaddis, J. 
Haugh, W. Heck, W. Heinrick, B. Hill, L. Hill, B. 
Johnson, J. Keith, L. Key, E. Knudtson, J. Kogan, 
C. Kowaleski, B. Lawhead, P. Lawson, J. 
Lindstrom, T. McEneaney, R. A. Moe, W. 
Netherton, J. Oar, D. O'Brien, R. Ogilvie, H. 
Postovit, R. Risebrough, D. Shields, P. Simonet, 

C. Stone, D. Struthers, S. Struthers, H. Tordoff, J. 
Trapp, L. Tynan, V. Wade, L. Walker (deceased), 

D. Whitcare, and S. Williams. 

The following agencies and organizations 
provided funding and logistical support for the 
surveys: U.S. Army Chemical Systems Labora- 
tory; the Danish Government; U.S. Air Force; 
World Wildlife Fund (Canada); La Ministere du 
Loisir, de la Chasse et de la Peche du Quebec; 
residents of Kuujjuaq and Tasiujaq, Quebec; 
Northwest Alaskan Pipeline Company (Flour 
Northwest Inc.); U.S. Fish and Wildlife Service; 
National Audubon Society; Province of Quebec 
Society for the Protection of Birds; Bell Museum 
of Natural History (University of Minnesota); 
Rocky Mountain Forest and Range Experiment 
Station, U.S. Forest Service; Bureau of Land 
Management (Arizona); Navajo Wildlife Branch 
(Arizona); Texas Parks and Wildlife Department; 
National Park Service; National Geographic 
Society; Fauna Silvestre of Mexico; British 
Columbia Fish and Wildlife Branch and the folks 
at North Island store (British Columbia). 

Literature Cited 

Ambrose, R. E., R. J. Ritchie, C. M. White, P. F. 
Scheitipf, T. Swen and R. Dittrick. 1988. Changes in 
status of Peregrine Falcon populations in Alaska. In 
Peregrine Falcon populations: their management and 
recovery. Edited by T. J. Cade, J. H. Enderson, C. G. 
Thelander and C. M. White. The Peregrines Fund, 
Inc., Boise, Idaho. 

Bird, D. M., and J. D. Weaver. 1988. Peregrine Falcon 
populations in IJngava Bay, Quebec, 1980-1985. In 
Peregrine Falcon populations: their management and 
recovery. Edited by T. .F. Cade, J. H. Enderson, C. G. 
Thelander and C. M. White. The Peregrine Fund, Inc., 
Boise, Idaho. 

Bromley, R. G. 1988. Statusof Peregrine Falcons in the 
Kitikmeot, Baffin, and Keewatin regions, Northwest 
Territories, 1982-1985. In Peregrine Falcon popula- 
tions: their management and recovery. Edited byT. J. 



1990 



White, Fyfe, and Lemon: 1980 Peregrine Falcon Survey 



181 



Cade, J. H. Enderson, C. G. Thelander and C. M. 
White. The Peregrine Fund Inc., Boise, Idaho. 

Bromley, R. G., and S. B. Matthews. 1988. Status of 
the Peregrine Falcon in the Mackenzie River Valley, 
Northwest Territories, 1969-1985. In Peregrine Falcon 
populations: their management and recovery. Edited 
by T. J. Cade, J. H. Enderson, C. G. Thelander and 
C. M. White. The Peregrine Fund, Inc., Boise, Idaho. 

Cade, T. J., and R. Fyfe. Editors. 1970. The North 
American Peregrine Survey, 1970. Canadian 
Field-Naturalist 84: 231-245. 

Cade, T. J., J. H. Enderson, C. G. Thelander, and C. M. 
VilhWt. Editors. 1988. Peregrine Falcon populations; 
their management and recovery. The Peregrine Fund, 
Inc., Boise, Idaho. 

Calef, G. W., and D. C. Heard. 1979. Reproductive 
success of Peregrine Falcons and other raptors at 
Wager Bay and Melville Peninsula, Northwest 
Territories. Auk 96: 662-674. 

Campbell, R. W., M. A. Paul, M. S. Rodway, and H. R. 
Carter. 1977. Tree-nesting Peregrine Falcons in 
British Columbia. Condor 79: 500-501. 

Ellis, D. H. 1988. Distribution, productivity and status 
of the Peregrine Falcon in Arizona. In Peregrine 
Falcon populations: their management and recovery. 
Edited by J. J. Cade, J. H. Enderson, C. G. Thelander 
and C. M. White. The Peregrine Fund, Inc., Boise, 
Idaho. 

Enderson, J. H., G. R. Craig, and W. A. Burnham. 1988. 
Status of Peregrines in the Rocky Mountains and 
Colorado Plateau. In Peregrine Falcon populations: 
their management and recovery. Edited by T. J. Cade, 
J. H. Enderson, C. G. Thelander and C. M. White. 
The Peregrine Fund, Inc., Boise, Idaho. 

Falk, K., and S. Moller. 1988. Status of the Peregrine 
Falcon in south Greenland: population density and 
reproduction. In Peregrine Falcon populations: their 
management and recovery. Edited by T. J. Cade, J. H. 
Enderson, C. G. Thelander and C. M. White. The 
Peregrine Fund, Inc., Boise, Idaho. 

Fyfe, R. W., S. A. Temple, and T. J. Cade. 1976. The 
1975 North American Peregrine Falcon survey. 
Canadian Field-Naturalist 90: 228-273. 

Hickey, J.J. Editor. 1969. Peregrine Falcon popula- 
tions: their biology and decline. Madison, University 
of Wisconsin Press. 



Hunt, W. G., J. H. Enderson, D. V. Lanning, M. A. 
Hitchcock, and B. S. Johnson. 1988. Nesting Pere- 
grines in Texas and Northern Mexico. In Peregrine 
Falcon populations: their management and recovery. 
Edited by T. J. Cade, J. H. Enderson, C. G. Thelander 
and C. M. White. The Peregrine Fund, Inc., Boise, 
Idaho. 

Kiff, L. F. 1988. Changes in the status of the Peregrine in 
North America: an overview. In Peregrine Falcon 
populations: their management and recovery. Edited by 
T. J. Cade, J. H. Enderson, C. G. Thelander and C. M. 
White. The Peregrine Fund, Inc., Boise, Idaho. 

Mattox, W. G., and W. S. Seegar. 1988. The Greenland 
Peregrine Falcon survey, 1972-1985, with emphasis on 
recent population status. In Peregrine Falcon 
populations: their management and recovery. Edited by 
T. J. Cade, J. H. Enderson, C. G. Thelander, and C. M. 
White. The Peregrine Fund, Inc., Boise, Idaho. 

Mossop, D. 1988. Current status of Peregrine Falcons in 
Yukon, Canada. In Peregrine Falcon populations: their 
management and recovery. Edited by T. J. Cade, J. H. 
Enderson, C. G. Thelander, and C. M. White. The 
Peregrine Fund, Inc., Boise, Idaho. 

Munro, W. T., and B. van Drimmelen. 1988. Status of 
Peregrines in the Queen Charlotte Islands, British 
Columbia, in Peregrine Falcon populations: their 
management and recovery. Edited by T. J. Cade, J. H. 
Enderson, C. G. Thelander, and C. M. White. The 
Peregrine Fund, Inc., Boise, Idaho. 

Porter, R. D., M. A. Jenkins, M. N. Kirven, D. W. 
Anderson, and J. O.Keith. 1988. Status and reproduc- 
tive performance of marine Peregrines in Baja 
California and the Gulf of California, Mexico. In 
Peregrine Falcon populations: their management and 
recovery. Edited by T. J. Cade, J. H. Enderson, C. G. 
Thelander, and C. M. White. The Peregrine Fund, Inc., 
Boise, Idaho. 

Walton, B. J., C. G. Thelander, and D. L. Harlow. 1988. 
The status of Peregrines nesting in California, Oregon, 
Washington, and Nevada. In Peregrine Falcon 
populations: their management and recovery. Edited by 
T. J. Cade, J. H. Enderson, C. G. Thelander, and C. M. 
White. The Peregrine Fund, Inc., Boise, Idaho. 

Received 8 June 1989 
Accepted 8 June 1989 



The 1985-1986 Canadian Peregrine Falcon, 
Falco peregrinus, Survey 



Julia E. Murphy 



Canadian Wildlife Service, Ottawa, Ontario KIA 0H3 

Present address: Faculty of Environmental Studies, York University, North York, Ontario M3J 1P3 

Murphy, Julia E. 1990. The 1985-1986 Canadian Peregrine Falcon, Falco peregrinus, survey. Canadian 
Field-Naturalist 104(2): 182-192. 

Results of Peregrine Falcon (Falco peregrinus) surveys carried out in 1985 and 1986 breeding seasons were used to 
assess the status of this species in Canada. Northern and west coast populations are stable or increasing. A new high 
number of nestings was reported in southern Canada. Comparison to the results of similar surveys in 1970, 1975 and 
1 980 shows the species is continuing to recover from the dramatic population decline it suffered prior to 1 970, although 
numbers remain low in southern Canada. 

Key Words: Peregrine Falcon, Falco peregrinus, populations, surveys. 

Les inventaires des populations de Faucons pelerins durant les saisons de reproduction 1985 et 1986 ont ete utilises 
pour etablir la situation de cette espece au Canada. Les populations du nord et de la cote ouest sont stables ou en 
croissance. Un nombre record de nids a ete enregistre au sud du Canada. Une comparaison avec les inventaires de 1970, 
1975 et 1980 montre que cette espece continue de se remettre du declin spectaculaire que les populations ont subi avant 
1970, bien que celles-ci restent faible dans le sud du Canada. 

Mots cles: Faucon pelerin, Falco peregrinus, population, inventaires. - 



Peregrine Falcon (Falco peregrinus) surveys 
were carried out across Canada during the 1985 
and 1986 breeding seasons. This report combines 
the results of these surveys to describe the status of 
this species in Canada. Similar surveys have been 
made at five-year intervals beginning in 1970 (Cade 
and Fyfe 1970; Fyfe et al. 1976; White et al. this 
issue). The 1970 survey documented severely 
reduced populations across Canada and Alaska 
and predicted possible extirpation of the species 
within the decade. The 1975 survey showed the 
Canadian population in continued decline. Results 
of the 1980 survey allowed guarded optimism; 
many population densities had improved and 
productivity was higher. In general, populations 
appeared to have stablilized, with decreases and 
increases on the local level. 

The results of the 1985 and 1986 surveys are 
encouraging. More nestings, especially by captive- 
released birds, were reported in the eastern part of 
the F. p. anatum range (Figure 1). than in any 
previous survey. F. p. anatum populations in the 
Northwest Territories and the Yukon Territory 
were stable or increasing, although 1986 was an 
unproductive year for the northern Alberta 
population. F. p. tundrius populations in the 
Northwest Territories appeared to be reproducing 
at normal rates, but the single F. p. tundrius 
population in Yukon Territory was extirpated. 
Because of the increasing frequency of observa- 
tions of captive-released birds, some information 
on captive-release programs in the provinces and 



the Yukon Territory is included here, although no 
attempt has been made to evaluate these programs. 

Eighteen areas in seven provinces and the two 
territories were surveyed by provincial and 
national governmental agencies and private 
groups. Three were surveyed in 1985, 13 in 1986 
and 2 in both years. Neither Manitoba nor 
Saskatchewan was surveyed but observations of 
peregrines occupying territories were recorded in 
both provinces. Because all areas were not 
surveyed in the same year, it is possible that birds 
moving from one survey area to another might 
have been counted twice or not at all. This is 
considered so rare as to be inconsequential because 
of nest-site fidelity, except where survey areas are 
very close together. 

Surveys across the range of Falco peregrinus 
anatum are reported here in detail because of the 
endangered status of this subspecies and the efforts 
invested in its reintroduction. Surveys of F. p. 
tundrius and F. p. pealei are described more 
briefly. 

Summaries of surveys conducted include 
information on survey methods, survey results, 
other noteworthy observations of peregrines in 
1985-1986, comparison with results of previous 
major surveys, and information on captive-release 
programs and population trends. 

Methods 

Breeding surveys of raptors are difficult. Several 
authors have described the errors, inherent biases, 



182 



1990 



MuRPHY: 1985-1986 Canadian Peregrine Falcon Survey 



183 



I ] TUNORIUS 

ANATUM 
B%^i PEALEI 




Figure 1. Known breeding distribution of the Peregrine Falcon in Canada (from Canadian anatum 
Peregrine Falcon Recovery Plan). 



lack of comparability, and other shortcomings 
(Brown 1974; Postupalsky 1974, 1981). Newton 
(1979) pointed out that raptors make poor subjects 
for study of population dynamics because they 
breed at low densities, often in remote and 



inaccessible places, and are frequently subject to 
human interference. 

In analysing and consolidating the data from the 
surveys, discrepancies in survey terminology, 
choice of population parameters, and methods for 



Table 1. Survey Terminology and Definitions. 



Term 



Definition 



Comment(s) 



Nest Site 

Occupied Nest Site or 
Territory 



Breeding Pair 
Productive Pair 



The actual site of the nest. 

A nest site or territory occupied by one 
or two territorial adults during all or 
some part of the breeding season. 



A pair which laid at least one egg. 

A pair which raised at least one chick to 
fledging, or, if actual fledging was not 
proven, raised at least one chick to an 
advanced stage of development from 
which the chick was assumed to have 
fledged. 



More than one alternate nest site may be 
present within a single territory. 

The terms active site and active territory 
have been used to describe such a diversity 
of situations that they have become 
meaningless and raptor biologists should 
avoid using them. 



For the purpose of Canadian surveys this 
term is more useful than the term Successful 
pair and its definition "a pair fledging at 
least one young", because it is often 
impossible to determine from infrequent 
aerial surveys whether nestlings fledge or 
not. C. Shank offered this definition based 
on the observation that nestling mortality is 
low from halfway through the nestling 
period to fledging. Brown (1974) identified 
the early part of the nestling period as one of 
the two times of strain in the breeding cycle 
of raptors. During this critical time only the 
male is available to obtain food for the 
young and the female. Later, food suply for 
the young is more assured because both 
parents hunt for the young and themselves. 



184 



The Canadian Field-Naturalist 



Vol. 104 



Table 2. Survey effort, 1985 or 1986, F. p. anatum. 







Number of 


Number 




Total 






historical 


of 




number of 




Year 


sites 


historical 


Other 


non-urban 




of 


known in 


sites 


areas/ sites 


sites 


Area 


survey 


survey areaf 


checked 


checked 


checked 


Labrador 


1985 


2 


2 


8 reported sites checked; other 
habitat surveyed incidentally 


10+ 


Bay of Fundy 


1986 


14 


14 


all cliffs along Bay of Fundy 


14+ 


Quebec 


1985 


10 


6* 


+ urban areas 


91* 


Ontario 


1985 


35 


35 


23 

+ urban areas 


58 


Manitoba 


no survey 








urban areas 





Saskatchewan 


no survey 


4 





urban areas 





Alberta South of 58° 


1986 


59 


59 


+ urban areas 


59 


North of 58° 


1986 


18 


16 





16 


Yukon Territory 


1986 










Porcupine River 




28 


10 


6 new occupied nest sites found 


10 


Yukon River 




30 


29 


1 new occupied nest site found 


29 


Peel River 




28 


13 


1 new occupied nest site found 


13 


Southern Lakes 




3 


3 





3 


Northwest Territories 












Mackenzie Valley 


1985 


84-89 


61 


total area checked = 4500 km^ 
17 new occupied nest sites found 


61 




1986 


101-106 


80 


total area checked = 4500 km^ 
5 new occupied nest sites found 


80 



fDoes not include sites found in survey year. 
*including Montreal 



their calculation, were encountered. The following 
discussion of these methodological concerns, as well 
as the issues of historical sites, non-breeding birds, 
timing of surveys and yearly population fluctua- 
tions, is intended to clarify the information 
presented in this report and contribute to the 
standardization of future surveys. 

Survey Terminology 

A major obstacle to interpretation of the survey 
data was inconsistent use of survey terminology. 
Various researchers used different terms to describe 
the same phenomenon; occasionally the same term 
was used to describe different phenomena in 
different survey reports. The terms and definitions 
in Table 1 have been reviewed by all surveyors and 
are used throughout this report. These terms should 
become standard terminology for future survey 
reports. 

Historical Sites 

As in previous surveys, particular attention was 
paid to historical nest sites as percent occupancy of 
historical sites is considered an indication of relative 
population size. Although this concept is valid, it is 
not equally applicable in all areas of Canada, as 
discussed below: 

1). In many areas historical records of peregrines 
are rare, not necessarily because the species was rare 
but possibly because it was not observed or 
observations were not recorded. For example, there 
are only 10 documented historical nest sites in 
Quebec south of Ungava Bay. 



2). The number of historical sites known is 
generally a total of sites used in many different years, 
including those used regularly and those used only 
infrequently, sites used only once in the recent past, 
perhaps even unsuccessfully, and sites that are no 
longer suitable nesting habitat because of 
development or disturbance. Long-term population 
records from Britain show that individual sites are 
not used with equal frequency. Ratcliffe (1980) 
categorized historical sites according to the average 
number of times used per decade. 

3). Peregrines are known to use alternate nest 
sites within the same territory. When only one of the 
alternate sites is known, surveyors may or may not 
record observations of use of another site within the 
same territory as occupation of a historical territory. 
This new site may or may not then be considered a 
separate historical site. 

4). Expanding and declining populations may 
behave differently from stable populations and from 
each other in their selection of nest sites. It could be 
misleading to assume that an expanding population 
will re-occupy traditional nest sites first. 

It is clear that the percentage occupancy figure 
should only be interpreted in the context of the 
extent of knowledge of historical populations in a 
given area, and caution should be used in comparing 
this figure for different areas. 

N on- Breeding Birds 

Observations of non-breeding birds (i.e. lone 
adults or sub-adults and non-breeding pairs) 



1990 



MuRPHY: 1985-1986 Canadian Peregrine Falcon Survey 



185 



Table 3. Occupancy in survey years 1985 or 1986, F. p. anatum. 







Total 






Occupancy 


Number 






number of 


Territories 


occupied by: 


of 


of 




Year 


territories 






historical 


urban 




of 


Occupied in 


a lone 




sites 


territories 


Area 


survey 


survey year 


adult 


a pair 


checked"* 


occupied 


Labrador 


1985 


2 





2 


1/2 (50%) 





Bay of Fundy 


1986 


1' 


1 


1 


1/14 (7%) 





Quebec 


1985 


1 





1 


1/6 (10%) 


1 


Ontario 


1985 


1 


1 





0/35 (0%) 


1 


Manitoba 


1986 


1 





1 


- 


1 


Saskatchewan 


1986 


2 


1 


P 


- 


1 


Alberta 


1986 












South of 58° 




2 





2 


0/59 (0%) 


2 


North of 58° 




6 


1 


5 


6/16 (38%) 





Yukon territory 


1985 












Porcupine River 




14 






8/10 (80%) 





Yukon River 




22 


N.D.3 




21/29 (72%) 





Peel River 




12 






11/13 (85%) 





Southern Lakes 











0/3 (0%) 





Northwest Territories 














Mackenzie Valley 


1985 


45 


N.D.3 




28/61 (46%) 







1986 


57 


N.D.3 




52/80 (65%) 






'Both lone adult and pair occupied same territory. 

^Male Peregrine Falcon mated with Female Prairie Falcon. 

^Not determined. Surveys could not determine whether sites were occupied by lone adults or pairs. 

''Does not include new sites found in survey year. 



Table 4. Reproduction in "remnant" F. p. anatum populations' in Northern Alberta, Yukon Territory and Northwest 
Territories'. 



Area 











Average no. 


Average no. 










young 


young 


Number 


Number 




Total young 


produced per 


produced 


of 


of 


Average 


produced by 


occupied 


per productive 


occupied 


productive 


clutch 


productive 


territory 


pair 


territories- 


pairs 


size 


pairs 


= productivity 




6 





3.8 
(n = 5) 











14 


11 


N.D.* 


29 


2.0 


2.6+ 1.0 


22 


18 


N.D. 


50 


2.2 


2.8 + 0.9 


12 



10 


N.D. 


23 


1.9 


2.3 + 0.8 


45 


36 


3.5 
(n=10) 


763 


1.7 


2.1 ±0.9 


57 


36 


2.9 
(n = 8) 


90" 


1.6 


2.5 + 0.9 



Alberta 

North of 58° 



1986 



Yukon Territory - 1986 

Porcupine River 

Yukon River 

Peel River 

Southern Lakes 
Northwest Territories 
Mackenzie Valley - 1985 

Mackenzie Valley - 1986 



*Not determined. 

'Does not include data on fostered and captive-reared young. 
^Includes territories occupied by lone adults and pairs. 

^Seven of 36 broods could not be counted. Figure given is mean brood size times number of productive nests. 
"•Six of 36 broods could not be counted. Figure given is mean brood size times number of productive nests. 



186 



The Canadian Field-Naturalist 



Vol. 104 



Table 5. Reproduction in "reintroduced" and Labrador F. p. anatum populations in survey and non-survey years' 





Number 


Number 


Number 


Number 


Number 


Number 




of lone 


of pairs 


of 


of eggs 


of 


of 




adults on 


occuying 


breeding 


in 


young 


young 


Area Year 


territories 


territories 


pairs 


nest 


in nest 


fledged 


Labrador 1985* 





2 


1 


4 


4 


3 


1986* 





2 


1 


2 


2 


? 


BayofFundy 1986* 


1 


I 





- 


- 


- 


Quebec 1985* 





1 


I 


32 





- 


1986 


1 


4 


3 


2 
1 
3 


2 


2 
















Ontario 1985* 


1 





- 


- 


- 


- 


1986 


1 


1 


1 


1 


1 


1 


Manitoba 1986 





1 





- 


- 


- 


Saskatchewan 1985 


1 


1 


P 


2 


2 


04 


1986 


1 


1 


P 


? 


? 


? 


Alberta 














South of 58° 1986* 





2 


2 


35 

46 


1 
3 


1 

3 


Young produced per occupied territory (productivity) (1985/86 average) = 


■■ 12/23 = .52 






Young produced per productive pair (1985/86 


average =12/ 


11 = 1.1. 









'indicates survey year 

'Does not include data on fostered young. 

^Female laid one egg in each of 3 different locations. 

-^Pair consisted of male Peregrine Falcon and female Prairie Falcon. 

"Hybrid young replaced by 3 captive-bred anatum young. 

^Edmonton 

^Calgary 



occupying nesting territories during the breeding 
season are very important. These instances of non- 
breeding are difficult to interpret as they may 
represent infertile birds or they may indicate that the 
population is expanding. For this reason, 
productivity calculated from all occupied territories 
is the most valuable parameter of reproductive 
success of a population (see below under 
"Parameters"). The proportion of territories 
occupied by lone birds is an indication of the size of 
the non-breeding adult population and may permit 
inferences about recruitment rates and the status of 
the population (Postupalsky 1974). 

Unfortunately, because lone adults and pairs 
without eggs are less attached to the nest site than 
are breeding pairs, they are more difficult to count 
accurately. Careful observations early in the season 
are required to find this component of the popula- 
tion and to determine that pairs have not attempted, 
unsuccessfully, to breed in a site unknown to the 
observer. Repeat visits may be necessary to confirm 
that the bird(s) is/ are actually occupying a territory. 
In the case of aerial surveys, when a single bird is 
seen on a territory, it is often impossible to 
determine whether the bird has a mate or not. 

Timing 

Fraser et al. (1983) and Postupalsky (1974) have 
described the biases introduced by surveys done at 



different stages of the breeding cycle. Surveys done 
at different times in different years are difficult to 
compare. 

Yearly Fluctuations 

Breeding populations of peregrines may show 
substantial yearly fluctuations (Court et al., this 
issue). The most obvious cause of these fluctuations 
is weather. For example, in 1986 a severe wind and 
rain storm in northern Alberta destroyed 9 of 19 
eggs and caused one nest to be abandoned; a late 
storm hit the Rankin Inlet area and many F. p. 
tundrius nests failed or were abandoned. Where 
populations are surveyed only once every five years, 
the exceptional nature of any one breeding season 
may not be immediately obvious. 

Parameters 

Brown (1974), Postupalsky (1974, 1981) and 
Fraser ( 1 978, in Postupalsky 1 98 1 ) all called for the 
standardization of population parameters and 
selected productivity as the most useful parameter of 
breeding success. Productivity is the average 
number of fledging or large young per occupied nest 
with known outcome, or the average number of 
young produced or recruited into the adult 
population per territorial pair. Previous surveys 
have reported this as "young per pair". As 
mentioned above, non-breeding pairs and single 



1990 



MuRPHY: 1985-1986 Canadian Peregrine Falcon Survey 



187 



Table 6. Results of Ungava Bay surveys 1970, 1975, 1980 and 1985. 











SL 


) Occupancy 


















Number 
















Total 




Historic 
















Historic 




Sites 








Lone 




Year 




Sites 




Checked 


Unoccupied 


Adult 


Pair 


1970' 




15 




15 




3 




3 


9 


1975' 




27 




25 




14 




2 


9 


1980' 




28 




21 




11 







10 


19852 




28 




20 




5 







23 

(8 new 

sites found) 


b) Reproduction 


















Young 


Young per 
















Total 


per pair 


pair with 






Pairs 


Pairs with 


Percent of 




young 


on 


young in 


Year 




Checked 


young 


total pairs 




in nests 


territory 


nest 


1970' 




9 


7 




78 




12 


1.3 


1.7 


19751 




9 


9 




100 




16 


1.8 


1.8 


1980' 




10 


10 




100 




27 


2.7 


2.7 


19852 




23 


19 




83 




61 


2.7 


3.2 


'White et al. 


(this issue). 
















2Lepage and Caron (1986) 

















birds assumed to have mates are included in this 
calculation and should be surveyed as accurately as 
possible. When the actual number of young in each 
nest known to be productive cannot be counted, an 
average figure for productive nests in which young 
were counted can be used. This introduces an 
element of error, but in the case of Yukon Territory 
and the Northwest Territories there is no alternative. 
Many raptor surveys also report young per 
successful pair and this has been done here. 

Results 

Survey effort is presented in Table 2, occupancy 
in Table 3, and reproduction in western and eastern 
Canada, respectively, in Tables 4 and 5. For two 
regions, Ungava Bay and Alberta south of 58°, 1985 
or 1986 survey results are presented along with 
historical survey data for 1970, 1975 and 1980 
(Tables 6 and 7). 



Regional Summaries 

Labrador Coast: — J. Brazil, Wildlife Division, 
Department of Culture, Recreation and Youth, 
P.O. Box 4750, St. John's, Newfoundland AlC 
5T7. 

1985 Survey: In 1985, 10 reported nest sites were 
visited between 2 July and 14 August. Two of these 
were sites where breeding pairs had been observed 
during the 1970 survey. Other potential habitat was 
investigated in the course of checking the ten sites 
(Figure 2). About 1 10 km of coastline was searched 
intensively by boat. A pair was found breeding at the 
historical site in Groswater Bay, Hamilton Inlet. On 
3 1 July the nest contained four downy chicks, one of 
which was dead (Table 5). No other peregrines were 
seen. Other observers also discovered one other pair 
and two single adults. The pair was observed 
regularly at Okak Bay, but no nest was located. 
Lone birds were found closer to Hamilton Inlet in 



Table 7. Survey results 1980-1986, Alberta, South of 58° 



Year 


Total Number 


Number 




Lone 






Known Territories 


checked 


Unoccupied 


Adults 


Pairs 


1970 


29 


29 


27 


1 


1 


1975 


35 


31 


30 


1 





1980 


53 


53 


53 








1986 


59 


59 


59 





2 urban' 



'Not included in 59 known territories. 



188 



The Canadian Field-Naturalist 



Vol. 104 



(^^ ON FOOT 

BY HELICOPTER 
^^ BY BOAT AND HELICOPTER 




Figure 2. Areas in Labrador surveyed in 1985. 



separate areas, but because they were only observed 
once during the breeding season, it was not possible 
to determine if they were actually occupying 
territories. 

1986 Survey: In 1986 the coastline between 
Cartwright and Makkovik was surveyed systemati- 
cally by boat and helicopter. The historical site in 
Groswater Bay was productive again; two young are 
believed to have fledged. Another pair was found 
occupying a territory at Cape Harrison on 28 July 
but no nest was found. 

A literature search has uncovered a number of 
additional historical records of peregrine nestings in 
Labrador, but none on the island of Newfoundland. 
These records date back as far as 1966 and seem to 
indicate that historically peregrines nested along 
many sections of the Labrador coast and possibly 
inland, not just in the areas surveyed in the early and 
mid-seventies. 

The Labrador coast was the last area in eastern 
Canada where peregrines are known to have nested 
prior to the beginning of captive-releases. During a 
1970 aerial survey of 200 km of coastline and 
offshore islands two nests with young were 
discovered, but the last known record of a 
productive nest in Labrador is from 1971. 
Continued investigations of nesting territories and 
reported observations up to 1980 failed to And any 
evidence of peregrines. 

The return of Peregrine Falcons to the Labrador 
coast between 1980 and 1985 is a hopeful sign, 
particularly since no releases have been made in 
Newfoundland and Labrador. It will be important 
to survey this area regularly in the future. 



Bay of Fundy: — B. Johnson, Canadian Wildlife 
Service, P.O. Box 1590, Sackville, New 
Brunswick EOA 3C0. 

On 1 July 1986 both the Nova Scotia and New 
Brunswick sides of the Bay of Fundy were surveyed. 
All known historical and potential nest cliffs were 
examined from a Cessna 337B aircraft. A reported 
inland historic site inland at Todd Mountain, York 
County, New Brunswick, was checked from an 
aircraft in late July. No peregrines or evidence of 
occupancy were found during these surveys. 

During 1986 a captive-release pair of adult 
peregrines occupied a territory near a hack site in 
Fundy National Park but did not nest. A lone male 
was also present in the same territory early in the 
summer. This may have been the sub-adult bird 
from a US captive release observed at the same site 
in 1985. 

Surveys carried out in the Maritime provinces in 
1970, 1975, and 1980, including a comprehensive 
survey of the Bay of Fundy in 1975 and a survey of 
the Nova Scotia side in 1980, failed to find any 
evidence of current or recent occupancy at any site. 

Quebec: — M. Lepage and M. Caron, Direction de 
la faune terrestre, Ministere du Loisir, de la 
Chasse et de la Peche, 150 boul. St-Cyrille est, 
Quebec, Quebec, GIR 4Y3. 

Southern Quebec: During the last week in June 
through the third week in July 1985 observers on 
foot or in helicopter examined, on one or two 
separate occasions, each of 91 potential nest sites 
within the area of southern Quebec shown in Figure 
3. Six of these were historical sites and 85 were 
potential sites identified by examination of 
topographical maps and aerial photos. 

The only occupied territory found in 1985 was in 
downtown Montreal. During the breeding season 
four different adults were present on this territory, 
including a female that laid three infertile eggs in 
three different locations. A team from the 
Macdonald Raptor Research Centre later inter- 
vened and provided two nestlings which the final 
pair raised to fledging. i 

In 1970 no evidence of peregrines was found at 
any historical sites, nor in two areas north of the St. 
Lawrence and east of Quebec City selected for 
examination because of their abundant cliffs and 
potential nesting sites. The conclusion then was that 
the peregrine had disappeared as a breeding bird 
from southern Quebec. Quebec was not surveyed 
again until 1980 when approximately 54 potential 
and historical sites in southern Quebec were 
surveyed and a successful nesting was documented 
at a historical site in the Eastern Townships. 

The attempted nesting in Montreal in 1985 was 
the eighth recorded over the six breeding seasons 
from 1980-85. A total of II wild young and 2 
fostered young were raised in these eight nests. In 



1990 



MuRPHY: 1985-1986 Canadian Peregrine Falcon Survey 



189 




Figure 3. Area of southern Quebec covered by the 1985 Survey. 



1986 four territories were occupied, three by pairs 
and one by a lone male. Two nests fledged a total of 
three young. 

Ungava Bay: It has not yet been established 
whether the peregrines nesting in the Ungava Bay 
area belong to F. p. tundrius or the F. p. anatum. 

In 1985 four areas in the Ungava Bay area were 
surveyed: the Gyrfalcon Islands; the Leaf River 
basin; the Payne-Arnaud River, from its mouth to 
the first rapids; and the Koksoak River, north of 
Kuujuaq. The Koksoak River was surveyed by 
canoe; all other areas were surveyed once by 
helicopter during the last week in July. 

Survey results are shown in Table 6 together 
withthe results of the 1970, 1975, and 1980 surveys. 

The areas surveyed in the Ungava Bay region 
have not been the same each time, so it could be 
misleading to compare the survey results directly. 
However, the population does seem to be 
reproducing effectively. Note that "young per pair 
with young in nest" may be an over-estimate of 
productivity because the chicks observed were 
quite young. 

Ontario: — Wildlife Branch, Ministry of Natural 

Resources, Whitney Block, Queen's Park, 

Toronto, Ontario M7A 1W3. 

Areas of eastern, central and northern Ontario 

were surveyed between early May and late July 

1985 (Figure 4). Eastern (more accessible) sites 

were reached by canoe; northern and central sites 

were reached by large boat or aircraft. Each site 

was visited once and whenever possible observed 



for five hours. Both historical (35) and potential 
(23) sites were visited. No evidence of nesting 
falcons was found at any of the 58 sites visited. A 
lone male occupied a territory in Arnprior. 

Surveys of all known historical sites and some 
potential sites in 1970 and 1980, and a partial 
survey of historical sites in 1975 failed to discover 
evidence of recent or current occupancy of any 
sites or non-breeding individuals. As in other 
provinces, it is possible that peregrines are nesting 
or have nested unobserved in small numbers in 
remote areas of the province. 

Although the results of the 1985 survey were not 
promising, encouraging finds were made in 
Ontario both before and after the year of the 
survey. In 1986 a successful nesting was 
documented at a non-urban site. It was not 
determined whether the adults were wild or 
captive-released birds. In 1983 a pair of peregrines 
nested in Arnprior. These birds had been released 
in Hull in 1 980 and 1981. Unfortunately the female 
was shot and killed and the two nestlings 
disappeared. The male returned to the site each 
year up to and including 1986 but has failed to 
attract a mate. 

Manitoba: — Wildlife Branch, Natural Resour- 
ces, Box 24, 1495 St. James St., Winnipeg, 
Manitoba R3H 0W9. 
No survey was carried out in Manitoba in either 

1985 or 1986. If Manitoba ever had a peregrine 

population it was probably very small. White et al. 

(this issue) were unaware of any confirmed nest 



190 



The Canadian Field-Naturalist 



Vol. 104 



y 

y 

1 

1 
1 
i 


> 

/ 
/ 
/ 


\ HUDSON V 
ONTARIO ^1 


1 

USA ^^' 




^ 1 


.r\ 


„ „.. 


J i^^'^'" 




' ' V 



Figure 4. AreasofOntariocoveredby the 1985 Survey. 



records for the province but Bechard (1981) 
reported a clutch of peregrine eggs collected near 
Gladstone, Manitoba, in 1887. No surveys have 
been conducted in Manitoba, with the exception of 
a brief unproductive survey in 1980 of an area of 
possible habitat along the lower Churchill River. 
In 1986, a pair of captive-released adult 
Peregrine Falcons was resident in downtown 
Winnipeg for more than six weeks and went 
through the motions of courtship, although no 
eggs were laid (Table 3). The female had been 
hacked in 1983 from the Montreal area, the male 
from Winnipeg in 1984. Unfortunately the male 
suffered an untimely death in a collision with a 
window or a wire. 

Saskatchewan: — Lynn Oliphant and Betsy 
Haug, Saskatchewan Cooperative Falcon 
Project, Department of Veterinary Anatomy, 
University of Saskatchewan, Saskatoon, 
Saskatchewan S7N OWO. 
No survey was carried out in Saskatchewan in 
1985 or 1986. There are few confirmed records of 
peregrine nests in the province. Neither a 1970 
survey of suitable habitat in southwestern 
Saskatchewan nor a 1 980 survey of a promising area 
north of Lake Athabasca turned up any evidence of 
breeding peregrines. However, recent sightings 
during the breeding season made by reliable obser- 
vers in the area surrounding Uranium City (Lake 
Athabasca) continue to suggest a small population 
in that area (L. Oliphant, personal communication). 
In 1985 a Saskatchewan rural-released male 
(1980) paired with a Prairie Falcon {Falco 
mexkanus) at a historical Prairie Falcon eyrie 
along the South Saskatchewan River (Table 3). 



Two hybrid young hatched and were removed and 
replaced by three captive-bred young peregrines, 
which were raised to fledging. In 1986 the young at 
this eyrie were found to be non-hybrid Prairie 
Falcons. A female peregrine was seen in the area 
several times. 

In April 1985 a lone adult male peregrine took up 
residence in downtown Saskatoon. He remained for 
the breeding season and fostered several captive- 
bred young to fledging. After returning in 1986 he 
was joined briefly by two females in sequence, each 
of which left before laying any eggs. He again 
successfully fostered several young. 

Alberta: — G. Erickson, Fish and Wildlife 

Division, Department of Forestry, Lands, and 

Wildlife, 9945-108 St., Edmonton, Alberta T5K 

2C9. 

Alberta was surveyed in 1986. As in past surveys 
(1975 and 1980) that part of the province south of 
58° was considered separately from the northern 
part. 

Alberta South of 58°: Between 16 and 19 May 
1986 a survey of all known historical sites (59) and 
other potential and reported sites was conducted 
by helicopter. No peregrines were observed. One 
hundred and thirty-one young captive-reared 
peregrines have been released over the period 1976- 
1985 in central and southern Alberta. Single pairs 
have established in the urban centres of Calgary 
and Edmonton, and all four adults are from the 
captive-release program (Table 5). The Edmonton 
pair established a territory in 1981 and nested in 
1982; the Calgary pair established a territory and 
nested in 1984. Pairs at both these locations 
successfully fledged their own young in 1986. 

Surveys documented the disappearance of 
breeding peregrines from natural habitat in 
southern Alberta and their re-appearance in urban 
centres (Table 7). Prior to the urban nestings the 
last confirmed successful nesting occurred in 1972 
(Fyfeetal. 1976). 

Alberta North of 58° : This area includes the only 
known remnant population of Peregrine Falcons 
in the prairie provinces. It has been intensively 
managed to maximize production since 1971 and 
surveyed annually since 1969. In no way can the 
results from this area be considered representative 
of a natural population. 

Over the period 1975-1986, 95 captive-raised 
young have been fostered to wild parents. Other 
manipulations have included double-clutching and 
the use of dummy eggs to keep pairs on sites until 
captive-reared young were available for fostering. 
Banded birds form a significant portion (55% in 
1986) of the adult population and captive-released 
birds breed regularly in the study area. 

In May and June of 1986, sixteen of the 18 
distinct peregrine territories previously identified 



1990 



MuRPHY: 1985-1986 Canadian Peregrine Falcon Survey 



191 



were ground-checked. (Two territories not 
surveyed are known to have been unoccupied over 
the period 1974 to 1985.) 

One territory was found occupied by a single male 
and five territories were found occupied by adult 
pairs (Table 3). All pairs laid eggs producing a total 
of 19 eggs. Five eggs were removed for pesticide 
analysis (one per nest), nine were lost or destroyed 
during a severe wind and rain storm in May and one 
nest was abandoned as a result of the storm. Only 
two eggs hatched, both from the same nest. Eight 
young from the captive-breeding facility at 
Wainwright were fostered to three of the pairs. Of 
the 10 young, only 3 survived to fledging. Four are 
believed to have succumbed to predators (including 
the two wild young), one was injured and subse- 
quently euthanized, and two died of unknown 
causes. 

British Columbia; — W. T. Munro, Wildlife 

Branch, Ministry of Environment and Parks, 

Victoria, British Columbia V8V 1X5. 

Coastal populations of Falco peregrinus pealei 

in British Columbia were investigated in the Queen 

Charlotte Islands, Gulf Islands, and northern 

Vancouver Island in 1986. Each area was surveyed 

once during the breeding season by boat and/ or 

helicopter; therefore only data on occupancy are 

available. Surveyors estimated at least 57 occupied 

territories in the Queen Charlotte Islands, four in 

the Gulf Islands, and nine on northern Vancouver 

Island (Checleset to Gordon Channel). 

Yukon Territory: — D. Mossop, Fish and 
Wildlife Branch, Dept. Renewable Resources, 
P.O. Box 2703, Whitehorse, Yukon Territory 
Y1A2C6. 

In the Yukon Territory there are five distinct 
subpopulations of Peregrine Falcons, each 
occupying different drainage basins separated by 
extensive mountain ranges. The subpopulation 
inhabiting the arctic coastal drainage is considered 
to be F. p. tundrius, and the four others, 
considered to be F. p. anatum, inhabit the 
Porcupine, Yukon, and Peel River drainages, and 
the nesting habitat associated with larger lakes in 
the southern portion of the territory. All of the 
subpopulations have been surveyed regularly since 
1973, although some have been investigated more 
frequently than others. Over the period 1973-1986, 
they have demonstrated different population 
dynamics. 

In 1986 each of the five populations was 
surveyed once or twice during the breeding season, 
either by helicopter or by boat (ground survey). 
The Yukon River population was surveyed from 
the ground at regular intervals during the breeding 
season. All or most of the known historical sites in 
each area were checked. 



Population Trends 

Porcupine River Drainage: The first evidence of a 
population recovery in the Yukon was reported in 
this area in 1981. Occupancy has remained high 
since then and productivity, though highly variable, 
is probably normal for these high-latitude anatum 
birds. 

Yukon River Drainage: Since 1980 this 
population has increased dramatically. In 1978 one 
nest was found, in 1986, 22 territories were occupied 
(Table 4). Active sites now outnumber formerly 
known historic sites almost 2:1. An equally striking 
increase has been documented in adjacent areas of 
Alaska. 

Peel River Drainage: This population has not 
been monitored as effectively as the above two. In 
the 1 980 and 1 98 1 surveys there was evidence of only 
a remnant group with very low productivity. In the 
1986 survey 72% of nesting sites checked produced 
young and two newly-occupied nesting territories 
were identified. This could represent an increasing 
trend in the population. 

Southern Lakes: This population is believed to be 
extirpated. It has been poorly known but visits to 
historic sites have failed to discover any evidence of 
occupancy. 

Arctic Coastal Drainage: (F. p. tundrius) Annual 
surveys since 1980 have covered most of the former 
range and all known historic sites. A dramatic 
decline has been reported. It is believed that the 
population ceased to breed in 1981, and no adults 
have been observed since then. 

Northwest Territories: — C. Shank, Wildlife 
Population Management, Department Renewa- 
ble Resources, Yellowknife, Northwest Territo- 
ries XI A 2L9. 

Both F. p. anatum and F. p. tundrius inhabit the 
Northwest Territories. The F.p. anatum population 
inhabits the Mackenzie Valley. 

Anatum Population: The Mackenzie Valley has 
been surveyed for Peregrine Falcons annually since 
1969, except 1983 (Bromley and Matthews, 1988). 
In 1985 and 1986, intensive helicopter surveys 
were made of the Mackenzie Valley between 
Norman Wells and Inuvik. In both years, a first 
survey was made in mid-June to document egg- 
laying followed by a second survey in mid-to-late 
July to quantify production. Each survey covered 
approximately 4 500 km^ checked most but not all 
historical sites. Survey effort was equivalent in both 
years. Seventeen new nest sites were found in 1985 
and five in 1986 bringing the total number known in 
the Mackenzie Valley to somewhere between 101 
and 106. 

Bromley and Matthews (1988) recently analysed 
the results of annual Mackenzie Valley peregrine 
surveys from 1969 to 1985. They concluded that the 
peregrine population declined by approximately 



192 



The Canadian Field-Naturalist 



Vol. 104 



35% in the late 1970s but returned to earlier levels by 
the mid-1980s. They found a parallel trend in the 
proportion of occupied sites where young were 
produced, but no observable trend in the mean 
number of young per productive nest. By 1985 the 
rate of population increase appeared to be declining 
or the population was leveUing off at a slightly lower 
level than during the early to mid-1970s. However, 
the 1986 occupancy of known sites (65%) and the 
calculated production of 90 young were higher than 
any previously recorded, which suggests the 
population may still be increasing. 

Tundrius Population: Bromley (1988) summar- 
ized the data available on populations in the 
Kitikmeot, Baffin, and Keewatin regions for the 
period 1982-1985. He concluded that all areas 
surveyed supported populations reproducing at 
normal rates. 

In 1986, tundrius peregrines were surveyed at 
Rankin Inlet (for the sixth year) and in the 
Kitikmeot region. Birds at both areas showed 
depressed reproductive success as a result of 
abnormally bad weather. 

Acknowledgments 

Appreciation is expressed for the invaluable 
contributions of the agencies and people who 
carried out the surveys and provided their results 
and feedback on earlier drafts of this document. 
T. C. Dauphine assisted with all stages of 
preparation of this report, I. M. Price and A. J. 
Erskine reviewed the manuscript. 

Literature Cited 

Bechard, M.J. 198L Historic nest records of the 
Peregrine Falcon in southern Saskatchewan and 
southern Manitoba. Blue Jay 89(3): 183. 

Bechard, M.J. 1982. Further evidence for a historic 
population of Peregrine Falcons in southern 
Saskatchewan. Blue Jay 40(2): 125. 

Bromley, R. C. 1988. Status of Peregrine Falcons (Fa/co 
peregrinus tundrius) in the Kitikmeot, Baffin, and 
Keewatin regions, Northwest Territories, 1982-1985. 
Pages 51-58 in Peregrine Falcon populations: their 
management and recovery. Edited kiyT. Cade, C. White, 
J. H. Enderson and C. G. Thelander. The Peregrine 
Fund. 

Bromley, R. G., and S. Matthews. 1988. Status of 
Peregrine Falcons {Faico peregrinus anatum) in the 



Mackenzie River Valley, Northwest Territories, 1969- 
1985. Pages 59-64 in Peregrine Falcon populations: their 
management and recovery. Edited byT. Cade, C. White, 
J. H. Enderson and C. G. Thelander. The Peregrine 
Fund. 

Brown, L. 1974. Data required for effective study of 
raptor populations. Pages 9-20 in Management of 
raptors. Edited by F. M. Hamerstrom, B. E. Harrell, 
and R. R. Olendorff. Raptor Research Report No. 2, 
Raptor Research Foundation, Vermillion, South 
Dakota. 146 pages. 

Cade, T. J., and R. W. Fyfe. 1970. The North American 
peregrine survey. Canadian Field-Naturalist 84: 231- 
245. 

Court, G. S., C. C. Gates, D. A. Boag, J. D. MacNeill, 
D. M. Bradley, A. C. Fesser, J. R. Patterson, G. B. 
Stenhouse, and L. W. Oliphant. 1990. A toxicological 
assessment of Peregrine Falcons, Falco peregrinus 
tundrius, breeding in the Keewatin District of the 
Northwest Territories, Canada. Canadian Field- 
Naturalist 104(2): 255-272. 

Fraser, J. D., L. D. Frenzel, J. E. Mathisen, F. Martin, 
and M. Shough. 1983. Scheduling Bald Eagle 
reproduction surveys. Wildlife Society Bulletin 11: 13- 
16. 

Fyfe, R. W., S. Temple, and T. J. Cade. 1976. The 1975 
North American Peregrine Falcon survey. Canadian 
Field-Naturalist 90: 228-273. 

Lepage, M., et M. Caron. 1986. Quatrieme inventaire 
quinquennal (1985) du faucon pelerin au Quebec. 
Ministere du Loisir, de la Chasse et de la Peche, Quebec. 
24 pages. 

Newton, L 1979. Population ecology of raptors. Buteo 
Books, Vermillion, South Dakota. 399 pages. 

Postupalsky, S. 1974. Raptor reproductive success: some 
problerhs with methods, criteria and terminology. Pages 
21-31 in Management of raptors. Edited by F. N. 
Hamerstrom, B. C. Hassel, and R. R. Olendorff. 
Raptor Research Report No. 2, Raptor Research 
Foundation, Vermillion, South Dakota. 146 pages. 

Postupalsky, S. 1 98 1 . Censusing nesting populations and 
measuring reproductive success. Pages 151-158 //? Eagle 
Valley Environmentalists Technical Report BED-81. 8 
pages. 

Ratcliffe, D. A. 1980. The Peregrine Falcon. Buteo 
Books, Vermillion, South Dakota. 416 pages. 

White, C. M., R. W. Fyfe, and D. Lemon. The 1980 
North American Peregrine Falcon survey. Canadian 
Field-Naturalist 104(2): 174-181. 

Received 22 February 1988 
Accepted 14 August 1989 



Status of the Peregrine Falcon, Falco peregrinus pealei, 

on Langara Island, Queen Charlotte Islands, British Columbia, 

1968-1989 



R. Wayne Nelson 

4218-63 Street, Camrose, Alberta T4V 2W2 

Nelson, R. Wayne. 1990. Status of the Peregrine Falcon, Falco peregrinus pealei, on Langara Island, Queen 
Charlotte Islands, British Columbia, 1968-1989. Canadian Field-Naturalist 104(2): 193-199. 

From 1968 to 1989, Langara Island, British Columbia, held 5-7 territorial pairs of Peregrine Falcons (Falco peregrinus 
pealei). Annualproductionfromtheisland varied greatly. In 1968-1979, it averaged 9.1 nestlings per year (range 5-13), 
and in the 1 980s it averaged 14.1 (range 8-20). Disappearance (presumed mortality) of known territorial birds averaged 
26% per year for males and 37% per year for females, higher than in five other studies of turnover in breeding 
Peregrines. Levels of DDE in falcon eggs in 1968-1972 were somewhat below the level that results in population 
decline. By 1986 the DDE had dropped to less than half of its former level. During the same period the PCB in eggs 
remained unchanged at moderate levels. The Langara Island nesting population of the Ancient Murrelet 
{Synthliboramphus antiquus), primary prey of the falcons, declined from very large numbers (probably well over 
250 000 pairs) in the 1950s and earlier, to 80 000-90 000 pairs in 1971, to 22 000-24 000 pairs in 1981 and 1988 surveys. 
Several possible causes of the murrelet decline are outlined, including changes in the ocean and predation by 
introduced rats. Between 1957 and 1968 the falcon population declined from 20+ pairs to 5 pairs and then remained at 
5-7 pairs through the 1970s and 1980s despite the further decline in the murrelet population. Although the number of 
pairs of falcons in earlier years appeared closely tied to the number of pairs of murrelets nesting on the island, from 
about 1968 onwards the Langara Island falcons apparently "relied" not on locally nesting murrelets but on murrelets 
commuting from distant nesting colonies. This falcon population in 1968-1989 was stable and reproductively healthy. 
A proposed rat eradication program might result in the recovery of the Langara Island murrelet population, and the 
falcon population, toward the very large numbers reported in the 1950s and earlier. 

De 1 968 a 1 989, il y avait sur Hie de Langara, Colombie-Britannique, de 5 a 7 couples territoriaux de Faucons pelerins 
(Falco peregrinus pealei). La production annuelle de Hie etait tres variable. Entre 1968 et 1979, il y eut une moyenne de 
9,1 oisillons par annee (entre 5 et 13), et pendant les annees 80, la moyenne fut de 14,1 (entre 8 et 20). Les disparitions 
(mortalite presumees) des oiseaux connus du territoire furent en moyenne de 26% par annee chez les males et de 37% 
par annee chez les femelles, chiffres plus eleves que dans cinq autres etudes sur la fluctuation du nombre des Faucons 
reproducteurs. Le niveau de DDE dans les oeufs entre 1968 et 1972 etait quelque peu en dela du niveau qui cause le 
declin des populations. En 1986, le taux de DZ)£'etait tombe a moins de la moitie du niveau anterieur. Durant la meme 
periode, les taux moderes de BPC dans les oeufs sont demeures constants. La population nicheuse d'Alques a cou 
blanc {Synthliboramphus antiquus) de Hie de Langara, la proie principale des faucons, tres nombreuse avant et 
pendants les annees 50 (probablement beaucoup plus de 250 000 couples), decUna a 80 000 ou 90 000 couples en 1971, 
et a 22 000 ou 24 000 couples aux releves de 1981 et de 1988. Plusieurs causes possibles du declin des alques ont ete 
evoquees dont des changements oceaniques et la predation par des rats importes. Entre 1957 et 1968, la population de 
faucons declina de plus de 20 couples a 5 couples et resta stable a 5 ou 7 couples pendant les annees 70 et 80 malgre le 
dechn continu de la population d'alques. Meme si, durant les annees precedantes, le nombre de couples de faucons 
semblait etroitement lie au nombre de couples d'alques nichant sur Hie, a partir de 1968 environ, les faucons de Hie de 
Langara ne furent plus "dependants" des alques locales, mais plutot d'alques de passage provenant de colonies 
distantes. Cette population de faucons, de 1968 a 1989, etait stable et se reproduisait bien. Un programme 
d'extermination des rats fut propose et pourrait bien etre la solution favorable au retabhssement de la population 
d'alques sur llle de Langara et ainsi ramener la population des faucons au nombre important releve pendant et avant 
les annees 50. 

Keywords: Peregrine Falcon, Falco peregrinus pealei. Ancient Murrelet, Synthliboramphus antiquus, predator-prey 
relationship, population dynamics. Queen Charlotte Islands. 

lht^tvQgnnQ.¥di\cons{Falco peregrinus pealei) upon small seabirds, primarily the Ancient 

nesting at Langara Island, British Columbia, have Murrelet {Synthliboramphus antiquus), these 

been visited by naturalists and scientists since 1915 falcons were only relatively lightly impacted 

(Green 1916) and recently have provided a during the 1950s- 1970s by the DDE-thin-eggshell 

continuous 22-year string of occupancy and syndrome that extirpated many other peregrine 

reproduction data, from 1968 to 1989. Because the populations (e.g. Hickey 1969, Cade et al. 1988). 

falcons occupying territories at Langara Island do The number of nesting pairs, reproductive success, 

not migrate, and because they prey almost entirely and a number of other aspects of the Langara 

193 



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Island falcons have been described earlier (e.g. 
Nelson 1970, 1976a, 1976b, 1977, 1980; Nelson in 
White et al. this issue; Nelson and Myres 1976). 
This paper reports on the status of this falcon 
population from 1968 through 1989, emphasizing 
the 1980s, summarizes recent Ancient Murrelet 
population changes, and re-evaluates the falcon- 
seabird relationship. 

Study Area and Methods 

Langara Island is at the northwestern tip of the 
Queen Charlotte Islands off the northern British 
Columbia coast. The island is about 10 km long 
and 6 km wide, heavily forested, with many chffs 
on its perimeter. 

During 1968, 1969, 1971, and 1972, my wife and 
I resided on Langara Island for 41/2 to 6 months 
spanning 15 February to 19 September and studied 
a number of aspects of the falcons' way of life. In 
1976, 1978, and 1979, basic occupancy and 
productivity data were obtained by the British 
Columbia Wildlife Branch. In each of the 
remaining years through 1989, with an assistant I 
visited the island usually for 8-10 days when 
nestlings were of banding age. The coastline cliffs 
were searched for territorial falcons by foot and 
from an inflatable boat. Although ocean 
conditions and time did not allow a thorough 
inspection of all of the coastal cliffs each year, a 
majority of the historical nesting sites and all 
recently occupied cliffs were checked each year. 
Usually the territorial falcons were conspicuous. If 
any territorial birds were missed over the years, the 
number must be very small. 

During the 1970s, usually the nest contents were 
observed only from a distance by telescope; in the 
1980s most nest cliffs were climbed, nestlings were 
banded and various data collected. Pop-rivetted 
green anodized aluminum color bands with unique 
letter and number combinations were placed on a 



majority of the nestlings in each year in 1984-1989; 
the bands were readable by telescope to about 100 m 
away. In 1968-1974 and 1980-1989, adult face 
markings were sketched or photographed through a 
telescope and used to identify the adults in 
subsequent years to determine turnover rates (see 
Nelson 1988). 

Background 

Beginning with Green (1916), Nelson and Myres 
(1976) summarized the ornithological literature 
describing the relatively large numbers of peregrines 
and the very large numbers of seabirds nesting on 
this island. The best-documented early report 
(Beebe 1960) listed 13 falcon nesting sites on the 
eastern half of the island that were used in most 
years during 1952-1958, and the whole island 
probably held 21-23 pairs of falcons at that time 
(Nelson and Myres 1976). The numbers of Ancient 
Murrelets in the 1950s and earlier, although not 
quantified by the various authors, were extremely 
large; in the early years the island almost certainly 
held more than 250 000 pairs and probably held 
between 375 000 and 750 000 pairs (Nelson 1977) 
and possibly was the largest Ancient Murrelet 
colony observed in this century. A dramatic drop in 
the number of murrelets nesting on the island 
apparently began in the mid-late 1950s and caused a 
very marked drop in the number of falcons nesting 
on the island by the time this study began in 1968. 

Concern for the welfare of the falcon population 
on the Queen Charlotte Islands as a whole prompted 
a review paper (Munro 1988) and a Commission of 
Inquiry (Shelford 1988), both of which pointed out 
the critical importance of the seabird populations in 
maintaining a strong falcon population. 

Breeding Population 

In each year from 1968 to 1979 the island held 
between 5 and 6'/4 territorial pairs of falcons (Nelson 



Table \. Occupancy and reproductive performance of Peregrine Falcons at Langara Island, British Columbia, 
during the 1980s. 

















Young per 




Number 


Lone 


Pairs with 


%of 


Total 


Young 


successful 


Year 


of pairs 


adult 


young 


total pairs 


young 


per pair 


pair 


1980 


6 





6 


100 


13 


2.17 


2.17 


1981 


6 





5 


83 


14 


2.33 


2.80 


1982 


6 





6 


100 


20 


3.33 


3.33 


1983 


7 





5 


71 


13 


1.86 


2.60 


1984 


7 





6 


86 


19 


2.71 


3.17 


1985 


5 


1 


4 


80 


8 


1.60 


2.00 


1986 


5 





4 


80 


11 


2.20 


2.75 


1987 


5 





4 


80 


13 


2.60 


3.25 


1988 


7 





6 


86 


17 


2.43 


2.83 


1989 


7 





5 


71 


13 


1.86 


2.60 


Average 


6.1 




5.1 


84 


14.1 


2.31 


2.76 



1990 



NELSON: The Peregrine Falcon on Langara Island 



195 



1988). During the 1980s the number of pairs 
fluctuated between 5 and 7 (Table 1). 

During 1970-1974, apparently at the end of the 
population decline for the falcons on this island, at 
three sites a situation termed pseudo-polyandry 
occurred in which a paired female prevented an 
adjacent, recently widowed male from obtaining a 
new mate (see Nelson and Myres 1976). At two of 
these sites, first one site (1972-1973) then the other 
site (1974) held a female that occupied both of the 
males' territories; by 1975 one of the males had 
disappeared and, through 1989, that part of the 
coastline then held only one pair of falcons. 

At another pair of sites, in 1972, the female at 
site C kept the neighboring male at site I single 
after his mate disappeared. However, when the site 
C female then disappeared, before spring 1973, 
both males obtained new females and then had 
offspring. In 1977-1978 site I was occupied; 
however, at site C in 1977 an adult female was 
briefly seen, but then drifted off, and in 1978 no 
falcons were found there. Both sites were occupied 
by pairs in 1979-1984. In 1985-1987 site I was 
vacant; in 1988-1989 both sites were again 
occupied by pairs. 

No instances of a single male on territory or a 
female exhibiting pseudo-polyandry were noted 
after 1974. That behavior appeared to be the means 
whereby the falcons enlarged their territories in 
response to a serious reduction in the numbers of 
their prey. 

In 1984-1985, a new pair was present at site A 
but produced no offspring. Only the female was 
found in 1986, and the site appeared vacant 
thereafter. Nearby, in 1988 at site D a new pair was 
found and that site produced nestlings in 1988 and 
1989. Both of these sites were historic sites that had 
not been active since the mid-1960s. 

The territorial behavior of these falcons has been 
investigated (Nelson 1977); they chase intruders to 
boundaries and fly display flights outward to, and 
along, territorial boundaries that are approxi- 
mately midway between neighboring nest cliffs. 
During this study, the territories of most of the 
pairs included two or more formerly distinct 
historic nesting territories. Occasionally, however, 
a new pair of falcons was able to become 
established and one territory was split into two. In 
1968-1989, the most stable situation appeared to be 
with 5 pairs, relatively widely spaced. The sixth site 
was occupied in a majority of years but in some 
years appeared to be incorporated within that of a 
neighboring pair. The seventh site (two nearby 
sites noted above) appeared to be more ephemeral 
and held a pair in only 4 years. This situation 
appeared to be normal. Under natural conditions 
peregrine populations elsewhere are remarkably 
stable and over a number of years show only minor 



fluctuations in the number of breeding pairs (often 
within 8% of the mean: Ratcliffe 1980:78). 

Productivity 

In a majority of years in the 1980s, only one pair 
failed to produce nestlings, twice two pairs had no 
nestUngs, and twice all pairs had nestlings (Table 
1). In 1968-1979 on the whole island the annual 
production averaged 9.1 nestlings (range 5-13). In 
the 1980s, however, the annual production from 
the whole island averaged 14.1 nestlings (range 8- 
20) and only twice was less than 13 nestlings. 
Production in 1982, 1984, and 1988 was especially 
good (Table 1). 

During the 1980s, the number of young per 
territorial pair was less than 2.0 during only 3 years 
in 10 (Table 1). During 1968-1979, however, there 
were less than 2.0 young per pair in 9 of 12 years. 
Similarly, during the 1980s the number of young 
per successful pair fell below 2.6 during 2 years in 
10, in contrast to falling below 2.6 during 9 of 12 
years in 1968-1979. It is suspected that weather 
factors were responsible for the larger brood sizes 
that were found at banding time in the 1980s, but 
statistical examinations have so far failed to detect 
any significant correlations between weather 
factor(s) and the average brood size or the total 
number of nestlings on the island in a given year. 

Survival of Breeders 

At Langara Island, territorial falcons that reared 
3-4 nestling (n = 44) disappeared from one year to 
the next at a rate of 43%, versus only 23% for 
territorial birds that reared only 0, 1, or 2 nestUng 
(n = 60) (0.90 < P < 0.95; Chi-square test). This 
suggests that when conditions result in a brood of 3 
or 4 nestlings the adult falcons provide optimal care 
for those nestUngs, absorb the additional costs of the 
larger brood, enter the following winter in poorer 
condition, and experience higher mortality rates 
(Nelson 1988). Overall annual loss of males was 26% 
and for females was 37% (n = 47 and 57 falcon-years 
respectively). These mortality rates were higher than 
any others recorded for peregrines (e.g. Mattox and 
Seegar 1988; Newton and Mearns 1988; Ambrose 
and Riddle 1988; Enderson and Craig 1988; Court et 
al. 1989) and probably resulted from the facts that 
the nesting falcons were resident year-round on the 
island and had to hunt seabirds over the ocean 
during very hostile winter conditions. Only in 1969 
and 1980 did a yearling female occupy a nest cliff on 
the island. In several years, over half of the adults 
recorded in one spring had disappeared and were 
replaced by the next spring, indicating that the 
productivity of this and nearby areas was good and 
that a substantial population of floaters (non- 
breeding adults) was available to replace lost 
breeders. 



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Vol. 104 



Biocides 

Nine falcon eggs (2 fresh, 7 addled) taken in 1968- 
1972 averaged, in parts per million (ppm) wet weight 
(range): DDE, 17.6 (10.9-25.7); and PCB, 11.0 
(3.24-35.5). The addled eggs were approximately 
25% lighter in weight (dehydrated) than the fresh 
eggs and so the organochlorines were somewhat 
concentrated. When "corrected" to fresh-laid 
moisture levels (Nelson and Myres, unpublished), 
the nine eggs averaged: DDE, 13.9 ppm (10.9-20.2); 
and PCB, 8.44 ppm (1.88 22.9). 

Eight of the eggs provided eggshell thickness 
indices which averaged 12.6% (4.2-19.9) less than 
pre-DDE eggshells from the Canadian west coast 
(pre- 1947 data of Anderson and Hickey 1972; 91 
eggs from Q.C.I., 80 of those from Langara). During 
1968-1972 a small number of falcon eggs at Langara 
Island disappeared during incubation, and a 
number of others failed to hatch, but the 
productivity was adequate to sustain the breeding 
population (Nelson and Myres 1976). DDE levels of 
15-20 ppm in Peregrine eggs are associated with 
reductions of 15-20% in the eggshell thickness index 
and the eggshell thickness itself. With more than 
that amount of eggshell thinning, too many eggs are 
broken, too few young are reared, and the 
population is unable to sustain itself (Fyfe et al. 
1988; Peakall and Kiff 1988). 

In 1986, four addled eggs from two nests 
averaged: DDE, 5.86 ppm (5.26 6.59); PCB, 8.58 
ppm (7.97-9.80); beta-BHC, 0.472 (0.424 0.559); 
heptachlor epoxide, 0.040 (0.037-0.043); and 
dieldrin, 0.028 (0.023-0.036). From 1968-1972 to 
1986, in falcon eggs the amount of DDE dropped to 
less than half and the amount of PCB remained the 
same. The 1986 levels of DDE were low enough that 
eggshell breakage would be a relatively rare event 
(see Nelson 1976b; Fyfe et al. 1988; Peakall and Kiff 
1988). 

Prey Base 

Earlier (Nelson 1976a, 1977, 1983) I suggested 
that (a) the stability fo the falcon breeding 
population at Langara Island since 1968 showed 
that the population of Ancient Murrelets on the 
island was also stable during this period, and (b) any 
further changes in this murrelet population would 
be reflected in parallel changes in the falcon 
population nesting on the island. Based on the 1971 
estimate by S. G. Sealy (in Vermeer et al. 1984) of 
80 000 90 000 pairs of Ancient Murrelets on the 
island, it appeared that each pair of falcons required 
between 13 000 and 22 000 pairs of murrelets to 
indefinitely sustain its harvest of roughly 1000 
murrelets annually, plus provide a protective buffer 
against overharvesting (Nelson 1977, 1983). 

Detailed surveys of the murrelet population in the 
1980s forced rethinking of details of this hypothesis. 



In 1981, Rodway et al. (1983), and Vermeer et al. 
(1984) used extensive searches, transects, and plots 
to locate the remaining colony areas, determine 
nesting burrow density, and estimate the total 
population of 22 500 pairs. In 1988 a more detailed 
survey was carried out, primarily in the main colony 
area located in 1981 (Bertram 1989). This survey 
estimated 24 000 pairs ± 4 000 (standard error) and 
showed that the colony occupied a smaller area but 
at a greater density of nesting burrows. Several lines 
of evidence suggested that the predation on 
murrelets by the long-established Alexandrian Rat 
{Rattus rattus alexandrinus) may have been an 
important factor in the population dechne of the 
murrelets (Bertram 1989). 

Although it seems clear that the murrelet decUne 
on the island between the mid-1950s and 1968 was 
responsible for a parallel decline in the falcon 
population, after 1968 the falcon population 
remained stable and did not parallel the further 
decline of the murrelets (by 60-75%) between 1968- 
1971 and the 1980s. Why did the falcon population 
not decline further? Were rats actually responsible 
for the murrelet decline in the 1950s- 1960s and more 
recently? 

Falcon-Seabird Relationships 

In the Aleutian Islands, Alaska, Amchitcka 
Island contained 16-22 occupied falcon territories in 
1968-1973 (White 1975). The nearest significant 
seabird colonies were on adjacent islands more than 
20 km away, and yet a majority of the diet of the 
nesting falcons was composed of seabirds, 
apparently taken in offshore hunting flights. The 
falcon nests tended to be closer together at the ends 
of the island, probably because seabirds tended to be 
funnelled past the ends of the island. A similar 
situation appears to have existed at Langara Island 
since some time just prior to 1968. At that time the 
reduced falcon population probably was capturing 
primarily offshore, commuting murrelets instead of 
locally nesting murrelets, and a further decline in the 
murrelets nesting on Langara Island would not 
cause a further decline in the falcon population. 

Frederick Island, 33 km south of Langara Island, 
in 1980 was estimated to be inhabited by 
68 400 ±8114 pairs of Ancient Murrelets and by 
89 850±I3 169 pairs of Cassin's Auklets 
(Ptychoramphus aleuticus) (Vermeer and Lemon 
1986). That island has held 3, and occasionally 4, 
pairs of falcons (W. Munro, personal communica- 
tion) and has potential nest cliffs and space to allow 
for at least six pairs if the food supply were more 
abundant and falcon territories consequently 
smaller (personal observations). 

Forrester Island and adjacent islets, Alaska, are 
65 km north of Langara Island, in 1969, 7 pairs of 
falcons were found here (V . Robards in White et al. 



1990 



NELSON: The Peregrine Falcon on Langara Island 



197 



1976) and in 1976 five pairs were seen (DeGange and 
Possardt in Sowls et al. 1978) although in 1976 the 
entire coastline was not thoroughly searched (T. 
DeGange, personal communication). In 1976 these 
islands were estimated to contain 33 800 pairs of 
Cassin's Auklets, 30 000 + pairs of Ancient 
Murrelets, 389 000 pairs of Fork-tailed and Leach's 
storm-petrels {Oceanodroma furcata and O. 
leucorhoa), 54 000 pairs of Rhinoceros Auklets 
{Cerorhinca monocerata), and 41 800 pairs of 
Tufted Puffins {Fratercula cirrhata) (Sowls et al. 
1978). The Rhinoceros Auklet may be rather heavy 
for a male Peregrine Falcon to carry and it probably 
is seldom captured by males when the other species 
are available. 

Each falcon requires about 200 g of prey per day, 
or approximately one Ancient Murrelet, one 
Cassin's Auklet, four storm-petrels, one-half a 
puffin, or one-half a Rhinoceros Auklet per day per 
falcon. On this basis, at Frederick Island in 1980 
each falcon territory represented approximately 
53 000 pairs of murrlets and auklets (ca. 21 200 kg). 
At Forrester Island, if we consider all six prey 
species and assume that the whole island complex 
held 7 pairs of falcons, each falcon territory in 1976 
represented approximately 50 400 pairs of "Ancient 
Murrelet equivalents" (ca. 20 160 kg). 

From these various lines of evidence it is clear that 
the earlier peregrine: murrelet ratio of 1 pair: 
13 000-22 000 pairs for Langara Island (based on 
the 1971 estimate of murrelet numbers) did not 
represent a sustainable predator-prey relationship, 
and the 5-7 pairs of Langara Island falcons at that 
time and through the 1970s and 1980s were 
primarily relying on murrelets from more distant 
colonies. From the foregoing it also is evident that, if 
the Ancient Murrelet were to disappear as a nesting 
species from Langara Island, there probably would 
remain 5-7 nesting pairs of falcons on that island, 
provided that the Frederick and Forrester Island 
seabird colonies remained relatively stable and 
continued to provide seabird prey that were 
available to the Langara Island falcons. 

Cause(s) of the Seabird and 
Falcon Declines 

Bertram (1989) found considerable evidence of 
rat predation upon adult Ancient Murrelets and 
their eggs on Langara Island in 1988 and suggested 
that rats may have caused the murrelet decline. 
Beebe (1953) may have been the first to report rats 
on the island, and he (1960) and other authors (e.g. 
Nelson and Myres 1976; Vermeer et al. 1984) 
suggested that the effect of the rats on the murrelet 
population was not especially serious. It appears to 
be unknown when rats became established here, 
perhaps with the very active Sea Otter trade in this 
area in the early 1800s, as a result of the 



establishment of Langara Lightstation in 1913, with 
the supplying of two radar stations on the island 
during World War II, or several commercial fishing 
camps in the 1940s and 1950s. It is not conclusively 
shown that rats were the primary factor in the 
murrelet and falcon declines. Could the murrelet 
population, ashore for only a few months each year, 
simply have swamped the rat population? One 
particular difficulty with the rat predation/ murrelet 
decline hypothesis is this: if rats brought the 
murrelet population from roughly 1 / 2 million pairs 
or more down to 80 000 90 000 pairs between, say, 
1955 and 1971, and from 80 000-90,000 pairs to 
about 22 500 pairs between 1971 and 1981, then why 
were the rats unable to seriously deplete or eliminate 
the murrelet population between 1981 and 1988? 

In the data of Beebe (1960; see Nelson and Myres 
1976) there was evidence that the falcon population 
entered serious dechne on Langara Island in 1958 
when 5 of 13 sites were unoccupied. Of interest is the 
fact that the number of pairs in the Glaucous- 
winged Gull {Larus glaucescens) colony and the 
Pelagic Cormorant {Phalacrocorax pelagicus) 
colony near Langara Lightstation in 1958-1959 
dropped to 1/2-1/3 of their previous numbers of 
nests (Drent and Guiguet 1961), and through the 
1980s those colonies remained at roughly 1/2 of 
their earlier size (unpublished data). This suggests 
that a relatively permanent change in the ocean- 
based food supply occurred about 1958-1959. In 
1957-1958 the NE Pacific Ocean warmed considera- 
bly, and this general area had warmer than normal 
waters into the 1970s (Nelson and Myres 1976). In 
the late 1970s and the 1980s the western Canadian 
ocean temperatures were particularly warm 
(Freeland 1990, personal communication). This 
change may have markedly affected food abun- 
dance or availability for murrelets, cormorants, and 
gulls. 

Another change, in the salmon stocks near 
Langara Island, occurred such that a thriving fishery 
was reduced through the 1960s and then seriously 
cut back during the 1970s. Long-time fishermen 
commented in the early 1970s regarding the much 
reduced abundance of surface "boils" of small fish 
versus earlier years. These "boils" often had salmon 
associated with them, and possibly were caused by 
the salmon attacking the schools of small fish and 
forcing them to the surface — where they were easily 
visible to and vulnerable to the various seabirds. 
Perhaps fewer salmon resulted in fewer surface 
"boils", reduced availability of food to the seabirds, 
and reduced numbers of nesting seabirds. 

Although it is recognized that rats on Langara 
Island are a significant but unquantified mortality 
factor for the Ancient Murrelets nesting there, the 
real role of the rats in the murrelet decline is not 
clear. There is also evidence that oceanic factors may 



198 



The Canadian Field-Naturalist 



Vol. 104 



have been responsible for the initial murrelet 
decUne, and indirectly for the falcon decline. 
Perhaps rat predation was then able to overwhelm 
the reduced murrelet population, and drive it 
downward. However, the apparent relative stability 
of the Langara Island murrelet population through 
the 1980s confuses the matter. 

The Canadian Wildlife Service and the British 
Columbia Wildlife Branch are investigating the 
feasibility of a rat eradication program on Langara 
Island (W. Munro, A. Edie, personal communica- 
tion). Such an experiment could be very difficult and 
costly, but it would be extremely valuable in 
clarifying the recent effects of the rats upon the 
murrelet population, and, if successful, it could 
allow the Ancient Murrelets, other seabirds, and 
Peregrine Falcons to rebuild to the truly spectacular 
numbers observed on Langara Island in the 1950s 
and eariier. 

Acknowledgments 

Many kind people and institutions have assisted 
with these studies. For financial and other assistance 
I am especially indebted to the British Columbia 
Wildlife Branch, Canadian Wildlife Service, 
Department of Biology at the University of Calgary, 
National Research Council of Canada (grants to 
M. T. Myres), the Frank M. Chapman Memorial 
Fund of the American Museum of Natural History, 
World Wildlife Fund (Canada), Queen Charlotte 
Islands Museum Society, Prince Rupert Fisher- 
men's Co-operative Association, Langara Fishing 
Lodge, and Transport Canada at Prince Rupert and 
Langara Island. I am especially thankful for helpful 
discussions and/ or help in the field provided by 
Frank L. Beebe, Donald A. Blood, J. Bristol Foster, 
Spencer G. Sealy, Michael Rodway, Douglas F. 
Bertram, Trisha and Nick Gessler, Bill Munro, Ben 
van Drimmelen, Colin Brookes, Morley Riske, 
Keith Hodson, Jay Page, Ervio Sian, Jennifer A. 
Nelson, David R. Pitt-Brooke, Richard W. Fyfe, M. 
Timothy Myres, and my wife, Alora L. Nelson. 

Literature Cited 

Ambrose, R. E., and K. E. Riddle. 1988. Population 
dispersal, turnover, and migration of Alaska Peregrines. 
Pages 677 684 in Peregrine Falcon populations: their 
management and recovery. Edited by T. J. Cade, J. H. 
Enderson, C. G. Thelander, and C. M. White. The 
Peregrine Fund, Boise, Idaho. 

Anderson, D. W., and J.J. Hickey. 1972. Eggshell 
changes in certain North American birds. Pages 5 14 -540 
in Proceedings of the XVth international Ornithological 
Congress. Edited by K. H. Voous. E. J. Brill, Leiden. 

Beebe, F. L. 195.3. The Peregrines of the northwest coast. 
Falconry News and Notes 1: 5 11. 

Beebe, F. L. I960. The marine Peregrines of the 
northwest Pacific coast. Condor 62: 145 189. 

Bertram, D. F. 1989. The status of Ancient Murrelets 
breeding on Langara Island, British Columbia, in 1988. 



Technical Report Series Number 59. Canadian Wildlife 
Service, Delta, British Columbia. 67 pages. 

Cade, T. J., J. H. Enderson, C. G. Thelander, and C. M. 
White. Editors. 1988. Peregrine Falcon populations: 
their management and recovery. The Peregrine Fund, 
Boise, Idaho. 949 pages. 

Cade, T. J., and R. Fyfe. Editors. 1970. The North 
American Peregrine survey, 1970. Canadian Field- 
Naturalist 84(3): 231-245. 

Court, G. S., D. M. Bradley, C. C. Gates, and D. A. Boag. 
1989. Turnover and recruitment in a tundra populatin 
of Peregrine Falcons Falco peregrinus. Ibis 131: 
487-496. 

Drent, R. H., and C.J. Guiguet. 1961. A catalogue of 
British Columbia sea-bird colonies. Ocasional Paper 
Number 12. British Columbia Provincial Museum, 
Victoria. 173 pages. 

Enderson, J. H., and G. R. Craig. 1988. Population 
turnover in Colorado Peregrines. Pages 685-688 in 
Peregrine Falcon populations: their management and 
recovery. Edited by T. J. Cade, J. H. Enderson, C. G. 
Thelander, and C. M. White. The Peregrine Fund, 
Boise, Idaho. 

Freeland, H. J. 1990. Sea surface temperatures along the 
coast of British Columbia: regional evidence for a 
warming trend. Canadian Journal of Fisheries and 
Aquatic Sciences 47: in press. 

Fyfe, R. W., R. W. Risebrough, J. G. Monk, W. M. 
Jarman, D. W. Anderson, L. F. Kiff, J. L. Lincer, 
I. C. T. Nisbet, W. Walker II, and B. J. Walton. 1988. 
DDE, productivity, and eggshell thickness relationships 
in the genus Falco. Pages 319 335 in Peregrine Falcon 
populations: their management and recovery. Edited by 
T. J. Cade, J. H. Enderson, C. G. Thelander, and C. M. 
White. The Peregrine Fund, Boise, Colorado. 

Fyfe, R. W., S. A. Temple, and T. J. Cade. Editors. 1976. 
The 1975 North American Peregrine Falcon survey. 
Canadian Field-Naturalist 90(3): 228-273. 

Green, C.deB. 1916. Note on the distribution and nesting 
habits of Falco peregrinus pealei Ridgeway. Ibis 58: 
473-476. 

Hickey, J. J. Editor. 1969. Peregrine Falcon popula- 
tions: their biology and decline. University of Wisconsin 
Press, Madison. 596 pages. 

Mattox, W. G., and W. S. Seegar. 1988. The Greenland 
Peregrine Falcon survey, 1972-1985, with emphasis on 
recent population status. Pages 27-36 in Peregrine 
Falcon populations: their management and recovery. 
Edited by T. J. Cade, J. H. Enderson, C. G. Thelander, 
and C. M. White. The Peregrine Fund, Boise, Idaho. 

Munro, W. T. 1988. Peale's Peregrine Falcon in British 
Columbia: status and management. Wildlife Branch, 
Victoria, British Columbia. 19 pages. 

Nelson, R. W. 1970. Langara Island, Queen Charlotte 
Islands. Pages 244 245 in The North American 
Peregrine survey, 1970. Edited bv T. J. Cade and R. 
Fyfe. Canadian Field Naturalist 84(3): 231 245. 

Nelson, R. W. 1976a. Langara Island, Queen Charlotte 
Islands. Pages 261 262 in The 1975 North American 
Peregrine Falcon survey. Edited by R. W. Fyfe, S. A. 
Temple, and T. J. Cade. Canadian Field-Naturalist 
90(3): 228 273. 

Nelson, R.W. 1976b. Behavioral aspects of egg breakage 
in Peregrine Falcons. Canadian Field Naturalist 90(3): 
320 329. 



1990 



NELSON: The Peregrine Falcon on Langara Island 



199 



Nelson, R. W. 1977. Behavioral ecology of coastal 
Peregrines {Falco peregrinus pealei). Ph.D. thesis, 
University of Calgary, Calgary, Alberta. 490 pages. 

Nelson, R. W. 1983. Natural regulation of raptor 
populations. Pages 126-150 in Symposium on natural 
regulation of wildlife populations. Edited by F. L. 
Bunnell, D. S. Eastman, and J. M. Peek. Proceedings 
Number 14, Forest, Wildlife and Range Experiment 
Station, University of Idaho, Moscow, Idaho. 

Nelson, R. W. 1988. Do large natural broods increase 
mortality of parent Peregrine Falcons? Pages 719-728 
in Peregrine Falcon populations: their management 
and recovery. Edited by T. J. Cade, J. H. Enderson, 
C. G. Thelander, and C. M. White. The Peregrine 
Fund, Boise, Idaho. 

Nelson, R. W., and M. T. Myres. 1976. Dechnes in 
populations of Peregrine Falcons and their seabird 
prey at Langara Island, British Columbia. Condor 78: 
281-293. 

Newton, L, and R. Mearns. 1988. Population ecology 
of Peregrines in south Scotland. Pages 651-665 in 
Peregrine Falcon populations: their management and 
recovery. Edited by T. J. Cade, J. H. Enderson, C. G. 
Thelander, and C. M. White. The Peregrine Fund, 
Boise, Idaho. 

Peakall, D. B., T.J. Cade, CM. White, and J. R. 
Haugh. 1975.. Organochlorine residues in Alaskan 
Peregrines. Pesticides Monitoring Journal 8: 255-260. 

Peakall, D. B., and L. F. Kiff. 1988. DDE contamina- 
tion in Peregrines and American Kestrels and its effect 
on reproduction. Pages 337-350 in Peregrine Falcon 
populations: their management and recovery. Edited 
by T. J. Cade, J. H. Enderson, C. G. Thelander, and 
C. M. White. The Peregrine Fund, Boise, Idaho. 

Ratcliffe, D. A. 1980. The peregrine falcon. Buteo 
Books, Vermillion, South Dakota. 416 pages. 



Rodway, M., N. Hillis, and L. Langley. 1983. Nesting 
population of Ancient Murrelets on Langara Island, 
British Columbia. Canadian Wildlife Service, Delta, 
British Columbia. Technical Report. 47 pages. 

Shelford, C. 1988. The falcon is telling us something. 
Report of the Committee of Inquiry on Falcons. 
Queen's Printer, Victoria, British Columbia. 60 pages. 

Sowls, A. L., S. A. Hatch, and C. J. Lensink. 1978. 
Catalog of Alaskan seabird colonies. United States 
Department of the Interior, FWS/OBS-78/78. 
Washington, D.C. 350 pages. 

Vermeer, K., and M. Lemon. 1986. Nesting habits and 
habitats of Ancient Murrelets and Cassin's Auklets in 
the Queen Charlotte Islands, British Columbia. 
Murrelet 67: 33-44. 

Vermeer, K., S. G. Sealy, M. Lemon, and M. Rodway. 
1984. Predation and potential environmental pertur- 
bances on Ancient Murrelets nesting in British 
Columbia. Pages 757-770 in Status and conservation of 
the world's seabirds. Edited by J. P. Croxall, P. G. H. 
Evans, and R. W. Schreiber. ICBP Technical Bulletin 
Number 2, Cambridge, England. 

White, C. M. 1975. Studies on Peregrine Falcons in the 
Aleutian Islands. Pages 33-50 in Population status of 
raptors. Edited by J. R. Murphy, C. M. White, and 
B. E. Harrell. Raptor Research Report Number 3. 

White, C. M., D. G. Roseneau, and M. Hehnke. 1976. 
Gulf of Alaska coast and southeastern Alaska. Pages 
259-261 in The 1975 North American Peregrine Falcon 
survey. Edited by R. W. Fyfe, S. A. Temple, and T. J. 
Cade. Canadian Field-Naturalist 90(3): 228-273. 

White, C. M., R. W. Fyfe, and D. B. Lemon. 1990. The 
1980 North American Peregrine Falcon, Falco 
peregrinus, survey. Canadian Field-Naturalist 104 (2): 
174-181. 

Received 5 June 1989 
Accepted 28 May 1990 



Preliminary Report on Breeding Peregrine Falcons, 

Falco peregrinus, in Labrador: 1987 and 1988 Survey Results 



D. Lemon' and J. Brazil2 



'Natural History Society of Newfoundland and Labrador, P.O. Box 1013, St. John's, Newfoundland AlC 5M3 
^Newfoundland Wildlife Division, Building 810, Pleasantville, St. John's, Newfoundland AlC 5T7 

Lemon, D., and J. Brazil. 1990. Preliminary report on breeding Peregrine Falcons, Falco peregrinus, in Labrador: 
1987 and 1988 survey results. Canadian Field-Naturalist 104(2): 200-202. 

Surveys were conducted in 1987 and 1988 to locate breeding Peregrine Falcons (Falco peregrinus) in Labrador. Active 
nests were located in a variety of habitats comprising coastal areas, including offshore islands and fiords, and interior 
river valleys. Seven active Peregrine Falcon nest sites have been located in Labrador. 

Key Words: Peregrine Falcon, Falco peregrinus, Labrador, active nests. 

Des etudes ont ete meneesen 1987 et 1988 pour repererdes Faucons pelerins nicheurs (Falco peregrinus) au Labrador. 
Des nids occupes ont ete localises dans plusieurs habitats qui comprennent la region cotiere, incluant les iles et les 
fjords elognes du littoral, ainsi que les cours d'eau a I'interieur des terres. Sept nids occupes de Faucons pelerins ont ete 
reperes au Labrador. 

Mots cles: Faucon pelerin, Falco peregrinus, Labrador, nidification. 



In response to national management and 
conservation efforts directed toward the recovery 
of Peregrine Falcon, Falco peregrinus, popula- 
tions in Canada, a need for surveys to locate 
breeding Peregrine Falcons in Labrador was 
identified. Prior to 1987, the Newfoundland 
Wildlife Division conducted small scale surveys 
along the portions of the coast and other surveys 
on an ad hoc basis to address this need (Joe Brazil, 
personal communication). In 1987 and 1988 the 
Natural History Society of Newfoundland and 
Labrador coordinated funding and logistics from a 
variety of government and non-government 
agencies enabling large scale, systematic helicopter 
surveys of the Labrador coastline and interior 
regions to be conducted. 

The baseline Peregrine Falcon population data 
obtained from these surveys represents the most 
current information on the Labrador population 
and should provide wildlife researchers and 
managers with valuable information on which to 
base conservation and management efforts. 

The area that has been surveyed by helicopter 
(Figure 1) is described in detail by Lopoukhine et 
al. (1977) and comprises primarily steep fiords, 
vast U-shaped river valleys and upland plateaus, 
steep cliffs exceeding 500 m in height and 
numerous offshore islands. Prior to the surveys, 
potential areas that might support nesting 
Peregrine Falcons were identified using 1 :250 000 
topographic maps. After commencing the 
surveys, the survey route chosen depended on the 
suitability of the immediate habitat. Survey 
routes were selected on a daily basis once the field 



work commenced. All surveys were conducted 
using a Bell 206L helicopter with three observers 
in addition to the pilot. Cliff areas that appeared 
to offer potential Peregrine Falcon nesting 
habitat where searched from the helicopter 
travelling at 70-100 km/h while trying to 
maintain a distance of approximately 50 m from 
the cliff. When an adult Peregrine Falcon was 
observed, an intensive search of the immediate 
and adjacent cliff area was conducted in an 
attempt to locate a nest site. 

In 1987 and 1988, 1204 km and 666 km of 
potential Peregrine Falcon nesting habitat was 
surveyed respectively (Figure 1). In 1987, three 
active Peregrine Falcon nests were located during 
this study and one additional nest site was located 
near Cartwright, Labrador during fieldwork in an 
unrelated study (Department of National Defence 
environmental research). The active nest site near 
Cartwright has been monitored since 1985 by the 
Newfoundland Wildlife Division following initial 
identification in the early 1970s (Cade and Fyfe 
1970). In 1988, three active Peregrine Falcon nests 
were located during this study. A total of seven 
active Peregrine Falcon nest sites have been 
located in Labrador in the past two years. Table I 
provides a detailed description of the seven nest 
sites located in Labrador during the past several 
years. 

Our 1987 and 1988 survey results are intended to 
provide current information on the baseline 
population estimates of the Peregrine Falcon in 
eastern Canada for regional and national 
management and conservation efforts. 



200 



1990 



Lemon and Brazil: Peregrine Falcons in Labrador 



201 




Figure 1. Areas in Labrador surveyed in 1987-1 



Acknowledgments 

We wish to thank the following agencies for 
financial and logistical contributions that made 
this study possible: World Wildlife Fund 
(Canada), Natural History Society of Newfound- 
land and Labrador, Newfoundland Wildlife 
Division, Canadian Wildlife Service (Atlantic 



Region), Department of National Defence, 
McNamara Construction Company (Division of 
George Wimpey Canada) and S. Fudge and 
Associates Ltd. R. I. Goudie was a member of the 
survey team in the 1987 and 1988 surveys. The 
superb skill of pilot G. Goodyear of Universal 
Helicopter (NFLD) Ltd. made the surveys a success. 



202 



The Canadian Field-Naturalist 



Vol. 104 



Table 1. Peregrine Falcon nest site characteristics in Labrador. 













Number 








Height above 


Height of 




Nest site 


of 


Age of 


Date of 


Nest 


ground/ water(m) 


cliff/ hill(m) 


Aspect 


area 


eyases 


eyases 


survey 


I 


800 


1000 


SSW 


lake 


3 


10 days 


7/31/87 


2 


150 


180 


E 


fiord 


2 


4-5 weeks 


7/31/87 


3 


140 


155 


SSE 


coastal island 


2 


3 weeks 


8/01/87 


4 


70 


80 


ESE 


coastal island 


1 


— 


7/23/87 


5 


40 


60 


S 


coast 


2 


1 -2 weeks 


7/24/88 


6 


50 


75 


S 


coastal island 


3 


1-2 weeks 


7/24/88 


7 


100 


140 


S 


river valley 


2 


1-2 weeks 


7/25/88 



Literature Cited 

Cade, T. J., and R. Fyfe, Editors. 1970. The North 
American Peregrine survey, 1970. Canadian 
Field-Naturalist 84: 231-245. 

Lopoukhine, N., N. A. Prout, and H. E. Hironen. 1977. 
The ecological land classification of Labrador: a 
reconnaissance. Ecological Land Classification Series, 
Number 4. Lands Directorate (Atlantic Region) 
Environment Management Service. Fisheries and 
Environment Canada. 85 pages plus map. 

Received 12 February 1989 
Accepted 12 February 1989 



Addenda 

The surveys conducted in 1987 and 1988 
continued in 1989 and completed a search for 
breeding Peregrine Falcons along the entire 
Labrador coast. Some additional work was 
conducted by the Newfoundland Wildlife Division 
and the Department of National Defence in 1989. A 
total of 26 breeding pairs of Peregrine Falcons are 
now known to occur on the Labrador coast 
representing a substantial population. 

David Lemon 

General Delivery, Topsail, CBS. Newfoundland 
AOA 3Y0. 



23 March 1990 



The Reintroduction of the Peregrine Falcon, 
Falco peregrinus anatum, into Southern Canada 

Geoffrey L. Holroyd^^ and Ursula Banaschi 



'Canadian Wildlife Service, 2nd Floor, 4999-98 Ave, Edmonton, Alberta T6B 2X3 
-Department of Forest Science, University of Alberta, Edmonton, Alberta T6G 2H1 

Holroyd, Geoffrey L. and Ursula Banasch. 1990. The reintroduction of the Peregrine Falcon, Falco peregrinus 
anatum, into southern Canada. Canadian Field-Naturalist 104(2): 203-208. 

The recovery of anatum Peregrine Falcon populations in southern Canada has depended upon the release of captive 
raised young. Between 1976 and 1987, 563 young falcons were released at 24 areas in southern Canada (an average of 
5.04 young per area per year; range 1 to 16). At a minimum, 35 (6.2%) peregrines returned at least one year after their 
release. Paired falcons were encountered significantly further from their release site than single falcons. Females 
returned further from their release sites than males but the difference was insignificant. Peregrines most often returned 
to the same kind of site (urban or rural) from which they were released. One half of the returning birds were first 
observed as single birds, at one year of age, and did not return. In 1988 there were 12 territorial pairs south of the boreal 
forest which produced 1 1 young. Recommendations to improve future release efforts are made. 

Le retablissement des populations de Faucons Pelerins anatum au Canada meridional depend de la remise en liberte de 
jeune eleves en captivite. Entre 1976 et 1987, 563 jeunes faucons au total ont ete remis en liberte dans 24 regions du 
Canada meridional (une moyennede 5,04jeunes par region par annee;valeursallantde 1 a 16). Au moins35(6,2%)des 
individus sont revenus au moins un an apres la mise en liberte. Les couples de faucons reviennent de fa9on plus loins 
des sites de lacher que les individus seuls. Les femelles reviennet plus loins des sites de lacher que les males, bien que la 
difference soit insignifiante. Les Faucons Pelerins reviennet plus vraisemblablement a des emplacements du meme 
type (urbains ou ruraux) que ceux dans lesquels ils ont ete remis liberte. La moitie des oiseaux qui reviennent ont 
d'abord ete observes seuls, a un age de un an, et ne reviennent que pour un an seulement. En 1988 il existait 12 couples 
territoriaux au sud de la foret boreale qui produisirent 1 1 jeunes. Des recommendations visant a ameliorer les futures 
remises en liberte sont presentees. 

Key Words: Peregrine Falcon, Falco peregrinus anatum, captive release, reintroduction, Canada. 



The decline of the Peregrine Falcon (Falco 
peregrinus) in the 1950s and 1960s in North 
America and Europe has been well documented 
(Hickey 1969). In Canada, by 1970, the anatum 
subspecies of the Peregrine Falcon was extirpated 
south of 60° N and east of the Rocky Mountains 
(Cade and Fyfe 1970) with the exception of 
unknown numbers in northern Alberta, near 
Ungava Bay, and along the Labrador coast. While 
peregrine populations increased in some parts of 
the eastern and western United States, the recovery 
in Canada has been slower (White, Fyfe and 
Lemon 1990; Murphy 1990). In 1970, in eastern 
North America, no known resident breeding 
populations existed south of the boreal forest 
(Cade and Fyfe 1970). The reestablishment of the 
peregrine therefore depended upon the release of 
captive raised falcons (Cade et al. 1988). 

Since 1975, government and non-government 
agencies released young anatum falcons into 
southern Canada using hacking and fostering 
techniques (Fyfe 1987, 1988). The goal of these 
releases is to establish self-sustaining populations of 
Peregrine Falcons in southern Canada east of the 
Rockies. The Anatum Peregrine Falcon Recovery 
Plan (Erickson et al. 1988) established nine 



management zones in the anatum range based on 
political and ecological boundaries (See Figure 1 in 
Murphy 1990). The objectives of the recovery 
program are to establish a minimum of 10 pairs per 
zone by 1992 and to have natural production of 15 
young per zone as a five year average by 1997. 

This plan also discusses the factors that limit the 
recovery of the anatum Peregrine Falcon in 
southern Canada and recommends a series of 
management actions in priority that are required to 
successfully recover these populations (Erickson et 
al. 1988). The release program is a priority 2 activity 
after priority I activities of monitoring the 
populations, protecting the populations from 
pesticides and preserving the anatum gene pool. The 
gene pool is protected through the captive breeding 
facilities of the Canadian Wildlife Service, 
Wainwright; University of Saskatchewan, Saska- 
toon; and Macdonald College, McGill University, 
Montreal (Fyfe 1988). The other factors and 
recommended actions are: priority 2 — preserve 
habitat; priority 3 — protect peregrines from human 
disturbance and predation, and mitigate the impacts 
of disease and disaster; and priority 4 — increase 
public awareness and conduct research and 
development to improve management techniques. 



203 



204 



The Canadian Field-Naturalist 



Vol. 104 



Table 1. Summary of releases of captive-raised Peregrine Falcons in southern Canada 





1976 


77 


78 


79 


80 


81 


82 


83 


84 


85 


86 


87 


Total 


southern Alberta 


11 


11 


6 


5 


23 


24 


22 


10 


11 


9 


- 


4 


136 


Saskatchewan 


- 


1 


2 


4 


8 


8 


4 


- 


- 


5 


- 


7 


39 


Manitoba 


- 


- 


- 


- 


- 


4 


4 


3 


7 


4 


4 


4 


30 


Ontario 


- 


4 


4 


8 


15 


16 


12 


3 


18 


14 


12 


24 


130 


Quebec 


10 


9 


8 


8 


13 


16 


7 


12 


26 


9 


20 


4 


142 


Maritimes 


- 


- 


- 


- 


- 


- 


5 


5 


16 


12 


21 


27 


86 


Total 


21 


25 


20 


25 


59 


68 


54 


33 


78 


53 


57 


70 


563 



This paper reviews the history of the releases and 
returns of captive raised Peregrine Falcons south 
of the boreal forest in Canada. The purpose is to 
evaluate some aspects of the release strategies and 
recommend alternatives to improve the success 
rate of the reintroductions. Not included here are 
foster releases in northern Alberta and Yukon 
Territories, as well as population surveys in 
Ungava Bay, the Labrador coast. Northwest 
Territories, Yukon Territories, and British 
Columbia. Foster releases occur when captive 
raised young are provided to territorial pairs of 
Peregrines to raise and fledge. 

Results 

Release Program 

Between 1975 and 1987, 563 captive raised 
Peregrine Falcons were released in southern 
Canada (Table 1). The sex ratio of the released 
young was relatively equal (49% male: 51% 
female). The first Canadian release occurred in 
northern Alberta in 1975, followed in 1976 by 
releases at four sites in southern and central 
Alberta and two sites in Quebec. Since then 
peregrines have been released in all provinces 
except British Columbia, Prince Edward Island 
and Newfoundland. All captive raised young in 
Canada were banded with a red anodized 
aluminum band displaying a vertical alpha- 
numeric code and a standard US Fish and Wildlife 
Service band on the other leg. 

There have been 1 10 releases. The largest releases 
were in Algonquin Provincial Park, Ontario (61 
young in 9 years); Hull/ Ottawa (55 young in 11 
years); Montreal (52 young in 12 years); and 
Kananaskis Country, Alberta (49 young in 3 years). 
Birds were released in 24 areas with an average of 
5.04 young per year per area. The number of young 
per release per year exceeded nine on 15 occasions. 

In 10 Canadian cities over II years, 264 young 
were released. The average number of young per 
area per year was 4.7 and exceeded nine on only 
three occasions. All urban releases were from 
buildings. 

Rural areas received 299 young for an average of 
5.6 young per area per year. These releases exceeded 



nine young per year on nine occasions. All rural 
releases were from cliffs in varied habitats. The 
habitat around the Algonquin and Kananaskis 
releases is heavily forested. There have been 82 
young released at six sites associated with prairie 
rivers and lakes of Alberta and Saskatchewan. 
Releases in Nova Scotia, New Brunswick and some 
in Quebec have been associated with marine 
shorelines. 

Most Canadian releases (95%) used the hacking 
technique where humans provided the food for the 
young until the young attained independence. At a 
few release sites, returning but non breeding falcons 
helped to feed the introduced young, e.g., 1987 at 
Fundy National Park (see also Barclay and Cade 
1983). At the Fundy site, the adult male also chased 
away Common Ravens {Corvus corax), potential 
predators that approached the release box (S. Hicks, 
personal communication). The extent of post- 
fledging training by these voluntary foster parents is 
unknown. 

The remaining 5% of captive raised young were 
fostered to breeding peregrines to augment their 
brood, to replace broken or cracked eggs, or pairs 
suspected of having laid thin-shelled eggs in the past. 
Young were used to supplement production at 
urban releases in Calgary, Edmonton, and 
Montreal. None of these 13 young is known to have 
returned. Young were given to Prairie Falcon (Fa/co 
mexicanus) pairs at three sites in southern Alberta 
but this was discontinued due to concern about 
interspecific imprinting (Bird et al. 1975). Again 
none of these 13 young is known to have returned. 

Peregrine Returns 

The exact number of released birds seen in 
Canada at least one year after their release is difficult 
to determine. Some birds are recorded at or near a 
release site, but their band status, hence their 
identity, remains unknown. At other times a red 
band, indicating a Canadian captive raised and 
released bird, is noted but the alpha-numeric code is 
not observed. Only when the code or the U.S. Fish 
and Wildlife Service band number is reported does 
the exact identity of the falcon and its history 
become known. 



1990 



HOLROYD AND BANASCH: REINTRODUCTION OF THE PEREGRINE FALCON 



205 



Table 2. Distance (km) between release site and return 
sighting of captive raised and released Peregrine Falcons 



Paired Falcons 
N Distance (+ SD) 



Single Falcons 
N Distance (+ SD) 



Males 
Females 



91 : 

283 



121 

;325 



33.3 + 90 
197.0+ 170 



The codes on the red bands were read on 35 
different falcons during the breeding seasons. 
Approximately 1 1 other falcons were observed 
with red bands but the codes were not read. In 
addition at least three birds were reported with 
only U.S. FWS bands; one wore a black band; 
another had a green band; and at least seven were 
unhanded. Overall approximately 55 falcons were 
observed in the breeding season and apparently on 
territory, between 1978 and 1989. An additional 19 
birds were reported but their band status was 
unknown. Some of these sightings could be repeats 
or falcons with bands that were observed in 
different years, thus obvious repeat records were 
removed from the totals presented here. 

The average distance between release and return 
sites for all returns is 130.3 km (S.D. 217.5) but is 
further for females (263.1 km, S.D. 292.4, range 
to 720) than for males (5 1.8 km, S.D. 101.8, range 
to 300); however, this difference is not significant 
(Mann-Whitney U test, P>0.1). Of the 35 
identified falcons, 16 of 22 males and 5 of 13 
females'returned to their release site. The distance 
between release and return sites is significantly 
greater for pairs than for singles (204.1 km versus 
60.6 km, Mann-Whitney U test, P<0.01). 
Eighteen falcons, 15 males and three females, were 
observed as single birds. The remaining 17 were 
observed with potential mates during the breeding 
season (Table 2). 

Hack sites and territories can be characterized as 
urban or rural. Most peregrines returned to a site 
similar to their release site (32 of 35, 9 1 %). This rate 
is lower in the USA (Cade and Bird 1990). 
However, young released at rural sites returned at 
a lower rate than those from urban sites. Of the 35 



identified falcons, 24 were released from urban 
sites representing 10% of releases at urban sites, 
while the other eleven (4%) were released from 
rural sites. These data could be biased since falcons 
that return to urban areas are presumably detected 
more easily. Five of the 1 1 rural releases were in the 
Bay of Fundy area where considerable effort was 
expended to locate territorial birds. At the two 
largest rural release sites, Algonquin Park and 
Kananaskis Country, falcons could nest unde- 
tected on many remote cliffs. Spring surveys are 
difficult in Algonquin because of late ice breakup 
(G. McKeating, personal communication). 
Alternately, many of the falcons that were released 
in forested areas may have died. Nevertheless, the 
fate of falcons released in forested areas remains 
unknown. 

Eighteen falcons, 13 males and 5 females, were 
first observed at one year of age while 10, 4 males 
and 6 females, were first observed at two years of 
age (Table 3). The oldest first time observation is of 
a single six-year-old male during 1987 in Winnipeg 
which was not reported in 1988. All observed 
females appeared by their third year. The average 
age of first appearance of males is 1.8 years, while 
for females it is 1 .9 (Table 3). Fifty percent of the 
falcons are seen for only one summer. Conse- 
quently the average age of last return is only 
slightly higher than that of first sightings (Table 3). 
There is a slight tendency for paired males to live 
longer on average than other categories, although 
two females were seven and eight years old in 1988. 

In 1980, peregrine pairs established territories 
and bred in southern Quebec (Bird and Aubry 
1982) and in Edmonton. These were the first 
recorded nesting by captive raised and hack 
released young in Canada. Since then, the number 
of known territorial pairs in southern Canada has 
increased to 12 in 1988 (Table 4). Quebec has the 
most pairs and singles. Recent releases in the 
Maritimes resulted in pairs but no young by 1988. 
At least six single birds were recorded in and near 
potential breeding sites in 1988. 

In 1980, a Quebec pair, both captive raised and 
released falcons, produced the first two young. The 



Table 3. Age of returning Peregrine Falcons at first and last sighting 







1 


2 


Age 


of return 

3 


sighting (years) 
4 5 


6 


7 


8 


mean S.D. 


Males: 


first 
last 
1988* 


13 
7 



4 
5 
1 




3 
1 



1 
1 

2 




2 



1 

2 











1 


1.78+ 1.35 
2.47+ 1.48 


Females: 


first 
last 
1988 


5 
2 
1 


6 
3 





2 
3 




1 
1 















1 





1 


1.88+ 1.13 
2.20 ± 1.43 



"If the last sighting was 1988, it is reported separately since these falcons may still be alive. 



206 



The Canadian Field-Naturalist 



Vol. 104 



Table 4. Number of territorial pairs and single Peregrine Falcons in 3 management zones as outlined in the Anatum 
Peregrine Falcon Recovery Plan 



1978 79 80 



82 



83 



84 



85 



87 



Zone 1 




southern 


pair 


Quebec 


single F 




unidentified 


Maritimes 


pair 




single M 




F 




unidentified 


Zone 2 




southern 


pair 


Ontario 


single M 




F 


Zone 3 




southern 


pair 


Alberta 


single M 




F 


Saskatchewan 


pair 




single M 


Manitoba 


pair 




single M 


Total birds 





2 
1 


1 
3 


3 


3 
1 

1 


6 

1 


1 


1 
1 


2 
2 


1 


1 
1 
1 


1 


2 


2 
1 


1 

22 


2 

1 
2 


2 


2 


2 
1 


2 
1 
1 


1 

2 
1 




1 


2 


1 


11 
1 

2 


11 


14 


21 


22 


35 



'Pair includes a falconry trained female that was introduced to a territorial male. 
^Females bred in the U.S. but were released in Ontario. 



number of young produced in subsequent years is 
difficult to calculate since eggs and captive raised 
young have been switched at some sites and usually 
only easily accessible urban sites were checked. 
Since 1980 12 wild young plus 14 captive raised 
young fledged in Edmonton and Calgary, while 27 
young fledged in Quebec including at least 1 1 in 
1988. 

Discussion 

The above returns pose several questions that 
require consideration in an evaluation of the 
success of the recovery effort. What are the origins 
of these returning birds? How many captive raised 
young were observed after at least one year? What 
characteristics of these releases might affect the 
chances of the falcons returning to Canada? In the 
conclusion we offer suggestions for future release 
efforts. 

Most territorial falcons (79%) originated from 
Canadian hack releases. One female, wearing a 
black band in Winnipeg during 1988, was from a 
1987 Minneapolis release (Tordoff and Redig 
1988; R. Nero, personal communication). Another 
female, that wore a blue band and bred in southern 
Quebec in 1980 was released in the north-eastern 
U.S. (Bird and Aubry 1982). Three falcons, 
wearing only U.S. FWS bands, were reported in 
southern Ontario and Quebec. While they were 
most likely banded as wild young in the United 



States, they could have been banded as passage 
birds at trapping stations. 

The distance between the release and return sites 
of falcons in Canada is comparable to those in the 
USA. Peregrines in the upper mid-western USA 
dispersed on average 117 km (Tordoff and Redig 
1988) compared to 130 km in Canada with females 
dispersing longer distances and more males 
returning to or near their release site. Likewise in 
the Rocky Mountains in the USA, females 
dispersed 279 km on average and males 68 km, 
similar to the Canadian distances of 263 km and 52 
km respectively (Burnham et al. 1988). Since the 
dispersal distance is large, particularly for females, 
the current emphasis for releases in Canada at 
potentially attractive nest sites may be unwar- 
ranted, especially if the site is inaccessible to 
humans or more costly to use than others. The 
abundance of single territorial males provides 
opportunities to release females that have b6en 
retrapped and held over the winter as was done in 
Saskatoon in 1988 (L. Oliphant, personal com- 
munication). This "manufactured" pair success- 
fully fledged fostered, captive raised young but 
none of their own. 

Six of seven unhanded peregrines, that were 
reported in southeastern Canada, were paired. The 
first sighting, a male, was paired with a red-banded 
female at Ste Foy, Quebec, in 1986. The second 
male appeared in 1987 at Pont Pierre Laporte, 



1990 HOLROYD AND BANASCH: REINTRODUCTION OF THE PEREGRINE FALCON 



207 



Quebec City, paired with a female that had been 
released in Toronto in 1984. The remaining five 
sightings occurred in 1988: a male in Montreal 
paired with a female wearing only an aluminum 
band; a male in Winnipeg paired with a 
Minneapolis released female; a pair in southern 
Ontario; and a female that overwintered in 
Ottawa. 

Unhanded falcons almost certainly are wild bred 
since all captive raised falcons released in Canada 
wear two bands that are unlikely to have been 
removed or lost. However, the origin of these 
unhanded falcons remains open to conjecture. 
They could be from wild sites in the USA as not all 
wild raised young were banded. However, five of 
seven unhanded falcons were males which tend to 
disperse short distances, i.e., an average of 56 km. 
This does present the possibility that some of these 
birds were raised at wild sites in Canada. Some of 
these sites are known but few young have been 
banded because of concern about disturbing the 
parents and young, the lack of qualified climbing 
banders, and the inaccessibility of some cliff sites, 
e.g., Saguenay River in Quebec. The first wild- 
raised Canadian falcons from captive raised and 
released falcons were two young at a southern 
Quebec site in 1980 (Bird and Aubry 1982). Thus 
these unhanded peregrines could be wild raised 
offspring of captive raised Canadian released 
falcons. 

The third possible source of these unhanded 
falcons is from unknown wild Canadian nests. 
Since the releases in Algonquin, Hull and 
Montreal began in 1976 and 1977, enough time 
passed for pairs to form and breed in the wild. This 
possibility implies the need to rigorously check 
remote cliffs in southern and central Ontario and 
Quebec in the five-year survey planned for 1990. 

Overall, relatively few Peregrine Falcons 
returned or were sighted in the breeding season. Of 
the 58 falcons with known band status, 46 (79%) 
had red bands and were from Canadian releases. If 
the same percent of unknown birds, 14 of 18, are 
also from Canadian releases, then 60 falcons 
(10.6%) returned from the 563 young released in 
southern Canada. 

In the Rocky Mountains of the USA, 673 young 
were released between 1976 and 1985 with a 
maximum of 20 returning birds in 1985 (Burnham 
et al. 1988), lower than the Canadian returns from 
slightly fewer releases. On the other hand, in the 
eastern USA between 1975 and 1981, 52 falcons 
(14.7%) returned from 353 released (Barclay and 
Cade 1983). Even higher are the return rates of 20% 
in all habitats between 1976 and 1979 and of 39% at 
coastal salt marshes in New Jersey (Barclay 1980). 

The return rate varies with location. Of the 
larger releases, the return rates were lowest at 



Kananaskis Country, Alberta and Algonquin 
Provincial Park, Ontario. However, young 
released at Algonquin could breed undetected in 
the remote forest lands of this region. This 
possibility is supported by the sighting of 
unhanded falcons in southern Ontario in 1988 and 
southern Quebec between 1986 and 1988. Three 
males returned to Kananaskis Country but no 
known pairs have formed. Nelson (1987:5) used 
survival rates of peregrines in Great Britain to 
argue that some of these released young should 
have survived and suggests "that the Peregrine 
recovery in Alberta is slowly occurring (unseen) in 
the eastern slopes of the Rocky Mountains . . .". 
He concluded that the major problem is the lack of 
effort invested to search for nesting peregrines in 
remote areas of the Rockies. 

Peregrine Falcons are most numerous in areas 
with high densities of suitable prey (Newton 1979; 
Ratcliffe 1980). High numbers of seabirds on 
coastal areas support breeding pairs (Nelson and 
Myres 1975; Ratcliffe 1980). Likewise tundra 
peregrines prey upon small mammals in arctic 
areas, especially when prey are in cyclical high 
numbers (Court et al. 1988). The abundance of 
available prey in forested areas as compared to 
open country habitats such as marshes affects the 
development of the peregrine's hunting behavior 
(Sherrod 1982). Cade and Bird (1990) reported 
that the majority of urban nest sites in eastern 
North America in 1980 were within 800 m of water. 
Barclay (1980:83) concluded "the local food 
resource may be one of the major determinants of 
the suitability of artificial release sites". 

The characteristics of the release site may be 
critical to the success of the recovery effort. 
Overall, urban releases resulted in proportionately 
more returns than rural releases. However, rural 
release sites around the Bay of Fundy with its tidal 
areas have had a higher return rate than other rural 
areas, consistent with the results from coastal New 
Jersey (Barclay 1980). Future release sites should 
have adequate prey to support inexperienced 
young Peregrine Falcons and returning birds. 

The recovery effort to reintroduce anatum 
Peregrine Falcons into southern Canada resulted 
in the release of 563 captive raised young between 
1975 and 1988, and in 1988 12 pairs existed where 
one occurred in 1970. However, the low return rate 
(10.6%) indicates some room for improvement in 
the release effort. We offer the following 
suggestions: 

(1) Larger releases, 20 to 30 young per area 
per year, will increase the Ukelihood of pairs 
becoming established; (2) the available prey 
base at all release sites must be abundant to 
increase the initial survival of fledgling 
falcons; (3) monitoring of potential breeding 



208 



The Canadian Field-Naturalist 



Vol. 104 



sites must be adequate to detect territorial 
pairs; (4) once territorial pairs are located, 
their band status should be determined to 
establish their origin, but only without undue 
disturbance; and (5) where lone territorial 
males occur, falconry trained females could be 
released to establish pairs and captive raised 
young introduced if the pair fails to breed. 

Acknowledgments 

We thank all those individuals and agencies that 
supported the Peregrine Falcon recovery effort 
over the past two decades. We particularly thank 
those who supplied us with resightings of 
peregrines in Canada including Bruce Johnson, 
Shaun Hicks, David Bird, Pierre Laporte, Michel 
LePage, Irene Bowman, Mary Ellen Foley, Bob 
Nero, Lynn Oliphant, Dale Hjertaas, Steve 
Brechtel, Dave Moore, Gary Erickson, and John 
Folinsbee. We thank Richard Fyfe, Harry 
Armbruster, Phil Trefry, Helen Trefry, Lynne 
Oliphant, Paddy Thompson, David Bird and Ian 
Ritchie who have done so much to provide young 
peregrines for release. The comments of David 
Bird, Roger Edwards, Gary Erickson, Bruce 
McGillivray, Gerry McKeating and David Peakall 
on earlier drafts of this manuscript are greatly 
appreciated. 

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Barclay, J. H. and T.J. Cade. 1983. Restoration of the 
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Bird, D. M., and Y. Aubry. 1982. Reproductive and 
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Bird, D. M., W. Burnham, and R. W. Fyfe. 1985. A 
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Burnham, W. A., W. Heinrich, C. Sandfort, E. Levine, 
D. O'Brien, and D.Konkel. 1988. Recovery effort for 
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1988. The population biology of Peregrine Falcons in 

the Keewatin district of the Northwest Territories, 

Canada. Pages 729-740 in Peregrine Falcon popula- 
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Mossop, B. Munro, R. Nero, C. Shank, and T. Weins. 

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by G. L. Holroyd, W. B. McGillivray, P. H. R. 

Stepney, D. M. Ealey, G. C. Trottier, and K. E. 

Eberhart. Provincial Museum of Alberta, Natural 

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Field-Naturalist 104(2): 182-192. 
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Peregrines? Alberta Naturalist 17: 4-9. 
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Peregrine populations and its seabird prey at Langara 

Island, British Columbia. Raptor Research Report 3: 

13-31. 
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A. D. Poyser, Berkhamstead, England. 
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Books, Vermillion, South Dakota. 
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Falcons after leaving the nest. Ph.D. dissertation, 

Cornell University, Ithaca, New York. 
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site selection, and age of first breeding in Peregrine 

Falcons released in the upper midwest. 1982-1988. Loon 

60: 148-151. 
White, C. M., and D. A. Boyce. 1988. An overview of 

Peregrine Falcon subspecies. Pages 789 812 in 

Peregrine Falcon populations: their management and 

recovery. Edited by T. J. Cade, .1. H. Enderson, C. G. 

Thelander and C. M. White. The Peregrine Fund Inc, 

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Received 8 .lune 1989 
Accepted 27 .lunc 1990 



Peregrine Falcons, Falco peregrinus. Nesting in an 
Urban Environment: A Review 



Tom J. Cade' and David M. Bird 

'The Peregrine Fund, World Center for Birds of Prey, 5666 West Flying Hawk Lane, Boise, Idaho 83709 
Macdonald Raptor Research Centre of McGill University, 21,111 Lakeshore Road, Ste. Anne de Bellevue, Quebec 
H9X ICO 

Cade, Tom J., and David M. Bird. 1990. Peregrine falcons, Falco peregrinus, nesting in an urban environment: a 
review. Canadian Field-Naturalist 104(2): 209-218. 

In 1988, 30 to 32 pairs of the Peregrine Falcon (Falco peregrinus) were present in at least 24 cities and towns in North 
America. Of 24 nesting pairs, only 3 failed and the remaining 21 pairs raised 42 to 45 fledglings (including some 
fostered young). The production of young per successful pair in 1988 was 2.45 for urban sites vs. 2.51 for rural sites. 
From 1975 to 1 988, 89 percent of 1 84 peregrines released at Canadian urban sites survived to the flying stage, and in the 
United States 83 percent of 52 urban-released (hacked) peregrines survived to independence. Bridge sites were 
associated with a low fledging success. Of 16 urban breeders of known origin, 7 came from rural sites. The height of 
buildings used ranged from 14 to 50 floors; east-facing sites were most preferred. Prey items were quite diverse with 
Rock Doves (Columba livid) being most common. The most significant hazards, especially to fledglings, were 
premature fledging, collisions, and poisoning from pest control programs. 

En 1 988, on a denombre de 30 a 32 couples de Faucons pelerins {Falco peregrinus) dans au moins 24 villes et villages en 
Amerique du nord. Sur un total de 24 couples reproducteurs, 21 couples ont eleve de 42 a 45 oisillons (dont quelques- 
uns provenaient de centres de reproduction) et trois couples seulement ont echoue. En 1988, le nombre moyen de 
jeunes par couple couveur etait de 2,45 pour les sites urbains, par comparaison avec 2,51 pour les sites ruraux. De 1 975 
a 1988, 89%des 184 Faucons pelerins relaches aux sites urbains canadiens ont survecujusqu'au stadede I'envol, et aux 
Etats-Unis 83% des 52 Faucons pelerins relaches aux sites urbaines ont survecujusqu'au stude de I'independance. Les 
lachers effectues a partir des ponts ont connu un taux faible de succes. Des 16 Faucons pelerins reproducteurs dont 
I'origine etait connue, sept provenaient de sites ruraux. La hauteur des immeubles utilises allait de 14 a 50 etages et les 
sites qui donnaient sur le cote est etaient les plus favorises. Tout en etant assez diverses, les proies les plus communes 
etaient les pigeons bisets {Columbia livia). Les risques les plus importants, particulierement en ce qui concerne les 
oisillons, etaient I'envol premature, les collisions et I'empoisonnement cause par les programmes de lutte contre les 
cuseaux nuisibles. 

Key Words: Peregrine Falcon, Falco peregrinus, breeding, reproduction, release, urban, cities, buildings. 

The Peregrine Falcon (Fa/co/7ere^n>7W5) seldom Doves {Columba livia) in Baltimore in 1940 and 

nested successfully in North American cities in the perhaps as early as 1930 (K. Carnie, personal 

era before the pesticides-induced population crash communication). 

of the 1950s. J. J. Hickey and D. W. Anderson The widespread extirpation of Peregrine 

attributed this failure "to a lack of the proper Falcons owing to the effects of DDT and dieldrin 

nesting substrate for their eggs, to the birds' noisy in the 1950s prevented any further exploration of 

intolerance of intrusion when they have young, urban habitats by falcons dispersing from the 

and to the absence of terrain in which the young countryside in search of new territories, 

can safely make their first flight" (Hickey 1969). Four decades later, in 1988, the situation is very 

There were several notable attempts in former different, as 30 to 32 pairs of peregrines were 

times: twice unsuccessfully owing to human present in at least 24 cities and towns spread across 

interference on the St. Regis Hotel in Manhattan the entire continent in both Canada and the United 

in the 1940s (Penny 1981), successfully on the old States, not to mention the frequent occurrence of 

City Hall tower of Philadelphia in 1946 and lower unmated falcons in these and other urban locales 

down in 1949(Groskin 1947, 1952), successfully on (Tables 1 and 2). This situation is not unique to 

a church steeple in Harrisburg, Pennsylvania North America, as peregrines have also taken up 

(Groskin 1952), and most successfully of all on the residence on buildings in Europe (Mebs 1988). 
Sun Life Assurance building in Montreal, Quebec, 

where the same female nested continually from Origins of Urban-nesting Peregrines 

1940 to 1952 and raised 22 young from 3 different This contemporary, urban population of 

males (Hall 1955). Several less certain cases have peregrines is largely composed of captively reared 

been rumored for the Los Angeles basin in the and released falcons from various restocking 

1940s. Peregrines were purported to hunt Rock projects in Canada and the United States. 

209 



210 



The Canadian Field-Naturalist 



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1990 



Cade and Bird: Peregine Falcons in an Urban Environment 



211 





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While many urban-nesting Peregrines were 
originally released in cities, some were not. For 
example, the 2 falcons that bred in Montreal in 
1984 had been released from cliff sites in Vermont 
and New York. Furthermore, none of the 4 pairs 
nesting in New York City include birds released 
there in earlier years. Of 9 birds released in Los 
Angeles in recent years, 5 became breeders at 2 nest 
sites in that city (B. Walton, personal communica- 
tion), but several subsequent replacements came 
from the countryside. 

The majority of 46 breeding peregrines in the 
eastern United States returned to the same type of 
site from which they fledged (Table 3). Of 16 urban 
breeders, 7 originated from rural sites, whereas the 
tower and cliff breeders were derived only from 
birds released at those types of sites. Twenty-nine 
of 32 peregrines released in Canada returned to a 
site similar to their release site (Holroyd and 
Banasch, this issue). 

It is also a common misconception that 
peregrines are only nesting in cities where releases 
have occurred. Nine of the 24 cities listed in Table 1 
that were home to pairs of falcons in 1988 have 
never had peregrines released in them. As their 
name suggests, these falcons are great wanderers 
and birds released in Toronto have ended up in 
Boston and Springfield. Montreal birds have 
dispersed to Winnipeg and Detroit, and a Detroit 
bird flew to Toronto, to name but a few cases. 

Although wild peregrines have nested from time 
to time on old European castles, cathedrals or 
other such structures (Mebs 1969; Saar 1988), it 
does appear that these captively reared and 
released peregrines have a stronger proclivity than 
wild birds to be attracted to man-made structures 
as nest sites and to enter urban environments. In 
addition to tall buildings, they have set up 
housekeeping on bridges, towers, the superstruc- 
ture of a decommissioned liberty ship, and in an 
old Raven's {Corvus corax) nest on top of a 
window-mounted air conditioner under the eaves 
of an abandoned hunting lodge. 

These reintroduced falcons may exhibit such 
plastic behavior because they are less afraid of 
human beings and their artifacts than wild birds are 
and because they also have a wider variety of early 
experiences with different, usually artificial, 
environmental situations than wild young do. 
Typically they hatch in incubators, are hand fed in 
the laboratory for a few days, than parent reared in a 
cage, fed again by humans in a hack box and during 
the fledgling period — all before they become 
independent, "wild" birds in the out-of-doors. 

Urban Releases of Peregrines 

Biologists first became intrigued by the idea of 
releasing peregrines in cities because of problems 



212 



The Canadian Field-Naturalist 



Vol. 104 



Table 2. Growth of the urban population of Peregrine Falcons in the eastern United States from 1980 to H 













Young per 




Breeding 


Productive 


Young 


Young per 


Successful 


Year 


Attempts 


Pairs 


Produced 


Attempt 


Attempt 


1980 


1 


1 


2 


2.0 


2.0 


1983 


3 


2 


5 


1.7 


2.5 


1984 


5 


4 


12 


2.4 


3.0 


1985 


5 


3 


11 


2.2 


3.7 


1986 


6 


5 


11 


1.8 


2.2 


1987 


11 


6 


14 


1.3 


2.3 


1988 


10 


8 


16 


1.6 


2.0 


Total 


41 


29 


71 


1.7 


2.5 



Data compiled by M. Gilroy. 



encountered in the countryside with natural 
predators of young falcons. It seemed obvious that 
metropolitan environments, at least at the 
skyscraper level, would be free of Raccoons 
{Procyon lotor). Red Foxes (Vulpes vulpes), and 
Great Horned Owls {Bubo virginianus), although 
an owl showed up in broad daylight on a high 
building to discourage a nesting attempt in 
Montreal in 1982! Also, cities abound in feral 
pigeons (generally regarded as the falcons' favorite 
food). Starlings {Sturnus vulgaris), and other 
suitable prey. Moreover, tall buildings mimic 
cliffs, and all that is needed to turn many 
skyscrapers into suitable eyries is to provide a box 
with sand and gravel to give the falcons an 
appropriate substrate in which to make their nest 
scrape, as was so successfully demonstrated by the 
Sun Life falcons in Montreal (Hall 1955). Yet 
another attractive feature is proximity to water, i.e. 
rivers, lakes and oceans. Of 19 urban nesting sites 
sampled from Table 1,12 were within 400 meters of 
water, 4 within 800 meters, and only 2 were not 
near any sizeable body of water. Presumably, the 
falcons associate such bodies of water with an 
abundance of waterbirds and perhaps with routes 
for migratory species. They also provide open 
hunting spaces for the falcons (Cade 1960). 

In cooperation with Dr. Heinz Meng, The 
Peregrine Fund carried out the first experimental 
release from a building in 1974, from the top of the 
Faculty Tower on the campus of the State 
University College at New Paltz, New York. The 
project has been described in detail by Kaufman 
and Meng (1975). 

Serious work on releasing peregrines from tall 
buildings began in 1976 when the Canadian 
Wildlife Service released birds in Edmonton and 
Montreal (Holroyd and Banasch, this issue). This 
program was later expanded to other Canadian 
cities, i.e. Calgary, Toronto, Ottawa-Hull, 
Winnipeg, Saskatoon, Quebec City, Stc. Catha- 
rines, and Brockviilc'. in 1979, The l*crcgrine Fund 



released young over a 2-year period from the roof 
of the Interior Department Building and from the 
old Smithsonian "castle" in Washington, D.C. In 
1980 they began releasing in Manhattan from the 
roof of the Metropolitan Life Building and, later, 
from the headquarters of Consolidated Edison 
Company of New York. 

Other U.S. urban release sites have included 
Portland (Maine), Norfolk, Philadelphia, Boston, 
Albany, Chicago, St. Louis, Detroit, Milwaukee, 
Minneapolis-St. Paul, Denver, Boise, and Los 
Angeles. 

Hazards to City-nesting Peregrines 

While all of the postulated advantages of urban 
environments proved true, there are also 
disadvantages mainly associated with life- 
threatening circumstances when the young falcons 
are developing their ability to fly. 

Deaths and serious injuries result when birds fly 
into plate glass windows and even into the sides of 
concrete buildings. Tinted and/ or mirrored glass 
buildings have proven particularly troublesome. 
Observers at 14 of 19 urban sites surveyed from 
Table 1 cited collisions with this type of building as 
their major concern. Often they result from some 
combination of inexperienced flight, strong and 
sometimes unexpected wind shears, and being 
startled either purposefully or inadvertently by 



Tabi.f. 3. Origins for known breeders in eastern United 
States Peregrine Falcon population. 

r, ,. Origin (hack or hatch site) 



Site 


Urban 


Tower 


Cliff 


Total 


Urban 


9 


3 


4 


16 


Tower 





23 





23 


Cliff 








7 


7 


lolal 


9 


26 


II 


46 



Data compiled by M. Gilroy. 



1990 



Cade and Bird: Peregine Falcons in an Urban Environment 



213 



human intrusion. Sudden gusts of wind have been 
known to sweep young falcons off nest ledges (J. 
Jennings, unpublished report). Paul Young 
(unpublished report) noted that some fledgUng 
peregrines lose altitude easily, i.e. 6 to 7 building 
floors per 10 sec of flight. He had to return several 
young birds to their nest site after grounding. 

Young peregrines have also been known to 
collide with wires and other artificial structures. 
Still other fledglings have lost their balance and 
fallen into open chimneys, air vents and even small 
courtyards with vertical walls, in which they 
became entrapped, or became entangled in fish line 
dangling from the girder of a building under 
construction. 

Several peregrines attracted to the open space 
and birdlife at nearby airports have collided with 
aircraft landing or taking off. One building used by 
peregrines had a heliport. While the loud noise and 
vibration seldom bothered the falcons, the female 
occasionally chased the departing helicopter for 30 
metres or so (P. Young, unpublished report). 

Perhaps the most serious problem facing young 
peregrines fledging in urban environments is their 
tendency to flutter or "helicopter" to street level 
(sometimes earlier than normal), usually unin- 
jured. This type of hazard occurs most often at 
buildings with sheer drop-offs to ground level, i.e. 
no terracing. Of 16 buildings housing peregrines in 
1988 (Table 1), 12 had such drop-offs. 

Once at the street level, the falcons are at risk 
from heavy traffic, dogs, people, and starvation. 
Often the adult birds are either reluctant to bring 
food down to grounded fledghngs or they cannot 
locate their young due to high noise levels and 
numerous nooks and crannies, e.g. parked 
vehicles, where the frightened birds can hide from 
view. 

Heavy traffic can be a problem for both young 
and adult birds. Falcons attempting to retrieve 
kills near busy freeways are at risk (P. Young, 
unpublished report). Young birds fluttering into 
heavy traffic have led to the appointment of traffic 
patrolmen to protect human rescuers, as well as 
posted warning signs in Salt Lake City in 1986 (W. 
Heinrich, personal communication). 

Bridges are particularly noteworthy for traffic 
fatalities among falcons. One of the Baltimore 
males was killed by a vehicle on the Francis Scott 
Key Bridge in Baltimore. Bridges have other 
problems, including a low fledging success rate. Of 
11 bridge sites in North America in 1988, only 5 
were successful in producing a total of 6 young 
(Table 4). Young birds often fledge into the water 
and may drown if not rescued by human observers. 
Other sites offer easy access to children with BB 
guns. Bridges with a lot of supporting structure, i.e. 
towers, piers, anchors and framework, offer more 



Table 4. Reproductive success of Peregrine Falcons 
nesting on bridges in North America in 1988. 





No. 


No. 


No. 


Location A 


ttempts 


Productive 


Young 


Walt Whitman Bridge, 










Philadelphia 








Throgs Neck Bridge, 







1* 


New York 








Commodore Barry ^ 


_ 


1 


1 


Bridge, Philadelphia 








Betsy Ross Bridge, 










Philadelphia 








Delaware Memorial 










Bridge West 








Verrazano Narrows 










Bridge, New York 








Girard Point Bridge, 




1 


2 


Philadelphia 








Francis Scott Key ] 


or 2? 


1 


1 


Bridge, Baltimore 








Oakland Bay Bridge, 




1 


1 


California 








Vincent Thomas Bridge, 










Long Beach, 








California 








Pierre Laporte Bridge, 1 




1 


1+ 


Quebec City 









*1 young fostered. 

Data compiled by M. Gilroy. 



landing and branching opportunities for both 
fledgling and adult birds as opposed to simple 
arching bridges. 

Toxicological Hazards 

Several peregrines in different cities, 3 of them in 
Baltimore, have died from eating strychnine- 
poisoned pigeons. Little is known of secondary 
poisoning effects from avitrol, particularly when 
used at higher than normal concentrations. Some 
suspect lead poisoning as a problem caused by 
pigeons feeding at street level. Higher lead 
concentrations have been reported in urban birds 
(Tansy and Roth 1970). More recently, blood 
samples from 20 adult Merlins {F. columbarius) 
feeding on House Sparrows {Passer domesticus) 
revealed potential lead problems (L. OHphant, 
personal communication). DDE and PCB levels in 
fledgling peregrines killed in collisions in Canadian 
cities in 1988 were deemed insignificant (Peakall et 
al., this issue). A more recent concern is the 
organophosphate, fenthion. There have been 
suspected cases of fenthion poisonings in 
Saskatoon MerUns. In a controlled laboratory 
study in 1988, K. Hunt, D. Bird and P. Mineau 
(unpublished data) fed House Sparrows exposed 
to fenthion to American Kestrels {F. sparverius) 
and all 14 falcons subsequently died. 



214 



The Canadian Field-Naturalist 



Vol. 104 



Table 5. Comparative production in urban vs. non- 
urban pairs of Peregrine Falcons breeding in the eastern 
United States, 1977-1988. 







Non- 






Urban 


Urban 


Total 


Breeding Attempts 


41 


134 


175 


Productive Pairs 


29 


103 


132 


Young Produced 


71 


258 


329 


Young per Attempt 


1.7 


1.9 


1.9 


Young per Successful 








Attempt 


2.5 


2.5 


2.5 



Data compiled by M. Gilroy. 



Other concerns include peregrines drinking or 
bathing in contaminated water collected in cooling 
units atop older buildings, or catching prey in 
sewage lagoons attracting myriad numbers of 
shorebirds. The effects of noise pollution from 
machinery like compressors and blowing fans is 
not known. Even human activity behind windows 
can disturb falcons on a ledge. For example, one 
pair which fledged 2 young in Los Angeles was 
composed of a male which often dozed only a 
meter away from workers in their offices. The 
female, on the other hand, vocalized at any 
excessive movement in the office and constantly 
slammed her feet into the window when food was 
brought to the young, whether people were present 
or not (P. Young, unpublished report). In contrast, 
another Los Angeles female was calm and 
approachable enough to allow Brian Walton to not 
only switch her eggs while she was sitting on them, 
but, on another occasion, even grab her to read the 
band numbers. Video cameras located less than 
five meters from the nest site have not disrupted 
nesting activities. 

Despite the varied and sometimes bizarre 
accidents that befall city-released peregrines, the 
success rate overall compares favorably with the 
results of release from towers in open country and 
from natural sites on cliffs. Jack Barclay of The 
Peregrine Fund analyzed the results of releasing 
peregrines in the eastern United States from 1975 
to 1981 and found that 83 percent of 52 falcons 
released at urban sites survived to independence. 
This figure compares well to birds released from 
towers (79 percent of 178) and to those released 
from cliffs (63 percent of 123). In more recent 
years, since moving release sites out of Great 
Horned Owl country, the latter figure has risen to 
80-85 percent and higher. Canadian releases 
between 1976 and 1985 were composed of 234 
Peregrines in rural sites and 184 in urban ones with 
88% and 89% of birds surviving to the flying stage, 
respectively (Fyfc, 1988). Hoiroyd and Banasch 
(this issue) reported that Canadian-released 



peregrines released at urban sites were more likely 
to return to nest than ones hacked from rural sites; 
however, they conceded that birds returning to 
urban sites may be easier to detect. 

Releases in cities have generally been successful, 
and they have contributed significantly to the 
establishment of an urban-nesting population, 
which is obviously expanding in size and 
distribution (Table 2). 

Reproductive Success 

In 1988, out of 24 nesting pairs, there were only 3 
failures (one of them fledged fostered young), and 
the 21 successful pairs (including those with 
fostered young) raised 42 to 45 fledglings (Table 1). 
From 1980 to 1988 in the eastern United States, 71 
percent of 41 nesting attempts in cities succeeded in 
producing an average of 1.7 young per attempt; at 
the same time, 77 percent of 134 attempts at non- 
urban sites produced an average of 1.9 young per 
attempt (Table 5). In short, the rate of 
reproduction in the urban birds compares well to 
the performance of healthy, expanding popula- 
tions nesting not only in the eastern United States, 
but also in Colorado (Enderson et al.), Scotland 
(Newton and Mearns), the French Jura Mountains 
(Monneret), and other reports in Cade et al. (1988). 

Other Concerns 

How many pairs of peregrines a given city or 
town can hold will, of course, depend on the its size 
and number of potential nest sites, e.g. buildings, 
bridges, etc. In Los Angelas, one pair nested on the 
Union Bank with another pair on the Cal Fed 
Building only 9 kilometres away. Currently in 
1989, there are 8 territorial pairs in the greater New 
York City region, and 2 nesting along the East 
River are only 3 kilometres apart. 

Releasing young peregrines in the vicinity of 
established nesting pairs or a single territorial male 
has often led to disaster, e.g. direct killing by adults 
or restricted access to food at hack sites leading to 
starvation. In some cases, a single adult male or a 
pair can be induced to become foster parents for 
young in hack boxes by playing taped recordings 
of begging calls (L. Oliphant, personal communi- 
cation). On the other hand, a male in Milwaukee in 
1988 killed 2 of his foster young after they fledged. 

Physical Features of Nest Sites 

Not all cities have environmental features that 
favor a high likelihood of success in either releasing 
birds or attracting a nesting pair. While a few pairs 
are nesting on fairly low buildings, including ones 
standing alone in smaller cities, the best results in 
releasing young (hacking) peregrines have been 
achieved, and most nesting pairs have been 
established, in cities with several tall buildings. 



1990 



Cade and Bird: Peregine Falcons in an Urban Environment 



215 



In a sample of 18 buildings chosen by nesting 
Peregrines in Table 1, 3 are the tallest in the city, 
and 11 are among the tallest. With one exception, 
all buildings were located only 1 block from other 
tall skyscrapers, and the proximity to water was 
noted earlier. 

Obviously, tall buildings are more visible to 
peregrines. Moreover, it is quite useful to have one 
or more surrounding buildings around the same 
height or slightly higher than the building used by 
the falcons or for release (hacking). It not only 
serves to give the fledged young a better view of the 
release box or nest, but can also act as an 
observation post for wardens. 

In a sample from Table 1, the average height on 
13 buildings where peregrines attempted to nest 
was 30 floors, ranging from 14 to 50 stories. The 
birds nested on average on the top 10% of the 
building, i.e. the 13th to the 50th floor. Five of 13 
pairs nested on the roof, while one pair on Hotel 
Utah in Salt Lake City raised 2 young in a nest tray 
on the 9th floor. 

Recommendations for Urban Releases 

When selecting sites for either release (hack) 
boxes or nest trays, it is advisable to stay away 
from rooftops or ledges that have dangerous 
machinery, open water air conditioning units, 
large fans, open shafts or chimneys, live wires, soft 
tar that might accumulate on their talons, etc. 
Certainly, restricting access to roof tops as 
opposed to ledges is inherently more difficult 
because of maintenance requirements. 

The architecture of old buildings like the Sun 
Life Assurance Building in Montreal provides 
numerous window ledges, cornices, or other 
convenient perches and potential nest sites at 
various levels. Descriptions of the physical features 
were obtained for as many of the building sites as 
possible listed in Table 1. One-half of 16 buildings 
used offered almost unlimited sites, 5 offered 2 to 4 
sites, while 3 had only 1 site. The fact that most 
buildings must be provided with gravel-filled nest 
trays to allow the falcons to make a nest begs the 
question as to how many trays should be provided, 
i.e. just one or several. Providing 4 sites on a 
Montreal building resulted in a clutch being split 
between 2 trays (D. Bird, unpublished data). 
According to T. French (personal communica- 
tion), a pair in Boston laid eggs in 4 different sites 
over two years. Even more prominent is the case of 
the peregrines laying 9 eggs in 9 different sites on a 
bridge in Quebec in 1988 (P. Laporte, unpublished 
data). 

Older style buildings with several small rooftops 
are more likely to offer a roof site without 
machinery and/ or heavy maintenance activity. 
Surprisingly though, but perhaps due to their 



greater availability, 13 "new" buildings as opposed 
to 7 "old" ones, i.e. over 20 years old, were chosen 
by nesting peregrines. As stated beforehand, 12 of 
16 buildings chosen had sheer drop-offs to street 
level. 

Ledge widths on 1 1 buildings chosen by 
peregrines ranged from a half-meter to 2 meters 
with a meter being most common. 

Despite the fact that 5 sites were biased because 
the nest boxes were deliberately placed facing east, 
that direction was most often chosen by nesting 
peregrines (10 sites). Other favoured directions 
were south (4), south-east (3), north (2), north-east 
(2), and 1 each for west, south-west and north- 
west. Presumably, a nest site protected from 
prevaiUng winds and excessive direct sunlight is 
preferred by peregrines. 

Clearly, managers fo both release sites and 
breeding pairs in urban environments can do much 
to minimize injury and mortality and enhance 
reproductive success. Buildings should be chosen 
where sheer, transparent glass facings on adjacent 
buildings are few or absent. There should also be 
few smoke stacks, air vents, air-conditioning units, 
or other mechanical contraptions which are likely 
to endanger recently fledged young, and all such 
hazards in the immediate environs should be 
screened from entry by peregrines. In Detroit, 
chimneys were capped and in Chicago, cooling 
units were covered with screening (P. Redig, 
personal communication) because one bird had 
previously drowned in such a unit. Ideally, there 
should be no major construction projects nearby. 
Discussions with pest control companies to curtail 
strychnine poisoning programs have been fruitful 
in some cities. The influence of sites brightly 
lighted at night on peregrine nesting success is not 
known, although some successful eyries such as the 
one in Baltimore receive illumination every night. 

Access to buildings on weekends and after work 
hours is generally necessary for peregrine 
management. Often, preliminary discussions with 
building owners and top level staff and mainte- 
nance personnel can circumvent many problems. 
In Los Angeles, for example, window washers 
refused to clean windows above the seventeenth 
floor on a building from March to June when 
Peregrines nested there. On that same building, 
maintenance workers were attacked and driven 
from the roof (P. Young, unpublished report). In 
Springfield, Massachusetts, a female peregrine 
defending its nest struck a biologist 8 times, even 
succeeding in knocking off his protective headgear 
which was strapped on (T. French, personal 
communication). Two other complaints from 
building owners included worries about liability 
and feathers from prey items clogging up 
ventilation screens. 



216 



The Canadian Field-Naturalist 



Vol. 104 



Not all urban-dwelling citizens want peregrines 
in the city either, especially pigeon fanciers. There 
are two sorts of pigeon fanciers who complain: 
those who like to feed feral pigeons in city parks, 
and the breeders of homing pigeons or other 
special varieties, such as tumblers and rollers 
which are especially vulnerable to hunting 
peregrines. The number of pigeon lofts in dense 
metropolitan living areas is sometimes unbelieva- 
ble, for example in the boroughs of New York City 
and in Los Angeles. Several reintroduced 
peregrines and their progeny have been shot in the 
Watts district of Los Angeles, where many inner 
city folk keep lofts contrary to city ordinance. No 
one has ever been arrested. For this reason the pair 
of peregrines nesting near this neighborhood is no 
longer allowed to fledge its young. The eggs or 
nestlings are removed and fostered elsewhere. In 
1989 however, a pigeon-fancier was arrested and 
fined for shooting a released captive-bred year-old 
peregrine in Sebastopol, California (A. Tennant, 
personal communication). 

Prey Taken by Urban Peregrines 

Previously, only Rock Doves and European 
Starlings {Sturnus vulgaris) were mentioned as 
food items of city peregrines by earlier authors 
(Culver 1919; Groskin 1947, 1952; Herbert and 
Herbert 1965). More recently. Barber and Barber 
(1983) reported the prey items taken over 3 years 
by Scarlett, the single female that inhabited 
downtown Baltimore from 1977 to 1983. Of 304 
prey items found by Barber and Barber, 277 (91 
percent) were Rock Doves. Certainly, pigeons do 
appear in the diet of peregrines in all of the 1 6 cities 
surveyed. In only one locale, Saskatoon, were 
pigeons considered unusual prey items for 
peregrines inhabiting that city (L. Oliphant, 
personal communication). However, pigeons were 
not necessarily the number one prey item in the 
other cities. For example, P. Redig (personal 
communication) noted that peregrines nesting in 
St. Paul and Rochester, Minnesota took many 
more pigeons than the pair in Minneapohs. 

Roller pigeons were particularly prevalent in the 
diet of Los Angeles peregrines (P. Young, 
unpublished report) because they are more 
noticeable in their peculiar flight pattern, fly at 
higher altitudes than feral types, and being smaller, 
can be more easily carried from a distance. Young 
also noted that most often pigeons taken were all 
or partially white, i-.e. brown and white, black and 
white. 

Eleven bird species caught during peak 
migration times and one mammal, i.e. Big Brown 
Bat i Eptesicus fuscus) comprised the remainder of 
prey items recorded by Barber and Barber (1983). 
While mammals arc somewhat unusual prey items 



for peregrines, bats are an exception (Cade 1982). 
The only other reports of mammals being taken 
were those of Norway Rats {Rattus norvegicus) 
eaten by peregrines nesting in a Montreal quarry in 
1983 (D. Bird, unpublished data) and ground 
squirrels {Spermophilus spp.) in Calgary (T. Nette, 
personal communication). Other noteworthy 
observations by the Barbers were the consumption 
of window-killed birds and extensive caching of 
food items. 

After Scarlett's death, the Barbers continued 
their study of the new pair of peregrines from 1 984 
to 1987 (Barber and Barber 1988). Their sample of 
472 items included 26 avian species, indicating a 
broad diversity for the Bakimore peregrines at 
least. This time. Rock Doves made up only 46% of 
all prey taken, whereas waterfowl and shorebirds 
comprised 22% of their prey. They suggested that 
the difference may be attributed to a different food 
preference of the new female, i.e. she spent her first 
15 months in the marshes of New Jersey prior to 
nesting in Baltimore. Having the smaller male 
present may also influence the diet of urban 
falcons. 

A seasonal trend was noted in the taking of 
pigeons. The Baltimore peregrines were less 
interested in catching pigeons during periods of 
high availability of either migrants or juvenile 
birds. In Boston, peregrines preyed heavily upon 
Starlings because of a major roost nearby (T. 
French, personal communication). 

Smaller raptors, i.e. American Kestrels {F. 
sparverius), Saw-whet Owls (Aegolius acadicus), 
have appeared in the diet of urban-dwelling 
peregrines. Two species noted by Barber and 
Barber (1983) for Baltimore were the Sora Rail 
{Porzana Carolina) and Yellow-billed Cuckoo 
{Coccyzus americanus). The Sora Rail, apparently 
easy to catch, has shown up with unexpectedly 
high frequency in Saskatoon (L. Oliphant, 
personal communication), Winnipeg (R. Nero, 
personal communication), Minneapohs (P. Redig, 
personal communication), and Edmonton (J. 
Folinsbee, personal communication). Yellow- 
billed Cuckoos were also taken in Saskatoon (L. 
Oliphant, personal communication) and Boston 
(T. French, personal communication), while 
Black-billed Cuckoos (C erythropthalmus) were 
killed by pairs in Boston, as well as in Minneapolis 
and Milwaukee (P. Redig, personal communica- 
tion). In Milwaukee, fledgling peregrines were 
hunting cuckoos after dusk and caching them. In 
other cities, the falcons have been seen flying 
during the night capturing migrants blinded by 
bright city lights (R. Thorsell, personal communi- 
cation). Perhaps not so surprising is the frequent 
appearance of domestic pet birds, i.e. various 
species of budgerigars {Melopsittacus), parakeets. 



1990 



Cade and Bird: Peregine Falcons in an Urban Environment 



217 



cockatiels and finches, either escaped or 
intentionally released. 

Paul Young (unpublished report) documented 
some of the hunting behaviour of Los Angeles 
peregrines. "Almost accipitrine in nature" using 
the element of surprise, most attacks were 
launched from a perch on a building with very little 
"waiting on", as peregrines typically do. The 
majority of hunting was done in the vicinity of the 
nest site. 

Attitudes Toward Urban Peregrines 

Like the pigeon fanciers mentioned earlier, there 
exist outspoken conservationists who decry the 
promotion of urban-nesting peregrines, some 
speaking from deep conviction and others from 
ignorance. Some naturalists feel that peregrines 
belong only on cliffs in wild and rugged country, 
and they object on esthetic and moral or religious 
grounds to deUberate attempts to establish falcons 
in man-created environments or to put them in 
places where they never occurred naturally. Some 
worry that having peregrines in cities might 
downplay the need to save habitat in the wild. 

Other critics have complained bitterly, some- 
times viciously, that urban releases are just 
elaborate publicity stunts by elitists or egomaniacs 
who want to attract attention to themselves or who 
want to raise money to support their enterprises. 
There is no question that great publicity attends 
the release of falcons in cities and, especially, the 
establishment of a successful breeding pair. 
Moreover, media people have at times been carried 
away in an excess of zeal to report on such events. 
Finally, it is understandable that the sponsors of 
city releases want to use the opportunity to 
advertise their business, as well as to raise funds 
"for conservation", not always for the support of 
further work on peregrines either. 

It is also undeniably true that city-dwelling 
peregrines provide unparallelled opportunities for 
public education and awareness about falcons and 
the problems of endangered wildlife. Urban 
peregrines have become the objects of much civic 
pride and enjoyment by millions of folk whose only 
other appreciation of wildlife comes from the 
occasional nature film on television. When 
"Scarlett" came to Baltimore and established 
herself on the 33rd floor of the United States 
Fidelity and Guaranty Building in 1977, she 
enlivened the whole city; and the history of her 
amours with "Rhet" and a series of mates over the 
next seven years captivated a following of many 
thousands of people across the nation. She died in 
1984 shortly after having raised a brood of her own 
young for the first time, but the USF&G eyrie 
remains occupied by a pair of falcons to this day; 
and there are few persons in the Baltimore area 



who could not give a fair account of peregrine 
biology and conservation. Much the same is true in 
other cities where peregrines are now nesting. 

Predictably, many local groups and state 
agencies have been overcome with "falcon fever" 
and now want to conduct their own city releases 
because of the favorable public relations and fund- 
raising opportunities associated with such 
activities. This enthusiasm is fine so long as the 
welfare of the peregrines and the priorities of the 
regional recovery plans are kept foremost. As long 
as the best interests and safety of the falcons 
remain the first concern, the secondary benefits of 
publicity, fund raising, and public education are 
added incentives, which no one should have to 
decry. Only when local groups or agencies begin to 
think first about publicity and money are things 
likely to go awry through a poor choice of location 
or birds for release. Those in charge of local 
projects must learn how to resist sponsors who 
have money but whose building is unsuitable. They 
should become wary of buying from commercial 
breeders, or entrepreneurial middlemen, falcons 
whose manner or rearing and whose geographic 
origins or genetic backgrounds are unknown or 
suspect, especially in Canada and the western 
states where it has always been official policy to 
release only birds of indigenous anatum stock. 

Based on the phenomenal increase in the 
population of urban-nesting peregrines and a 
reproductive success parallelling that of rural- 
nesting birds, it is safe to say that urban peregrines 
are here to stay and to multiply! There is obviously 
much interchange between urban and rural sites 
and hence, the pairs occupying urban sites should 
be viewed as integral parts of the overall 
population and not as oddities. 

Acknowledgments 

We are extremely grateful to the following 
people and the many organizations they represent: 
U. Banasch, J. Barber, M. Barber, J. Barclay, M. 
Byrd, K. Carnie, J. Carreiro, J. Folinsbee, T. 
French, R. W. Fyfe, R. Galbraith, M. Gilroy, W. 
Heinrich, J. Jennings, P. Laporte, M. Lepage, T. 
Nette, R. Nero, L. Oliphant, P. Redig, L Ritchie, 
G. Septon, R. Smith, M. Spreyer, H. Tordoff, B. 
Walton, J. Weaver, and P. Young. We also thank 
the Raptor Research Foundation, Inc. for 
allowing us to conduct a workshop on urban 
peregrines at the 1988 meeting in Minneapolis. 

Literature Cited 

Barber, J., and M. Barber. 1983. Prey of an urban 
peregrine falcon. Maryland Birdlife 39: 108-110. 

Barber, J., and M. Barber. 1988. Prey of an urban 
peregrine falcon — Part II. Maryland Birdlife 44: 
37-39. 



218 



The Canadian Field-Naturalist 



Vol. 104 



Cade, T.J. 1960. Ecology of the Peregrine and 
Gyrfalcon populations in Alaska. University of 
California Publications in Zoology 63: 151-290. 

Cade, T. J. 1982. The falcons of the world. William 
Collins Sons, London. 

Cade, T. J., J. H. Enderson, C. G. Thelander, and C. M. 
White. Editors. 1988. Peregrine Falcon populations: 
their management and recovery. The Peregrine Fund, 
Inc. Boise, Idaho. 

Culver, D. E. 1919. Duck hawks wintering in the center 
of Philadelphia. Auk 36: 108-109. 

Fyfe, R. W. 1988. The Canadian peregrine falcon 
recovery program, 1967-1985. Pages 599-610 in 
Peregrine Falcon populations: their management and 
recovery. Edited by T. J. Cade, J. H. Enderson, C. G. 
Thelander and C. M. White. The Peregrine Fund, Inc., 
Boise, Idaho. 

Groskin, H. 1947. Duck hawks breeding in the business 
center of Philadelphia, Pennsylvania. Auk 64: 
312-314. 

Groskin, H. 1952. Observations of duck hawks nesting 
on man-made structures. Auk 69: 246-253. 

Hall, G. A. 1955. Great moments in action: the story of 
the Sun Life Falcons. Mercury Press, Montreal. 

Herbert, R. A., and K. G. S. Herbert. 1965. Behavior 
of Peregrine Falcons in the New York City region. Auk 
82: 62-94. 

Hickey, J.J. Editor. 1969. Peregrine falcon popula- 
tions: their biology and decline. University of 
Wisconsin Press, Madison. 



Holroyd, G. L., and U. Banasch. 1990. The reintroduc- 
tion of the Peregrine Falcon, Falco peregrinus anatum 
into southern Canada. Canadian Field-Naturalist 
104(2): 203-208. 

Kaufman, J., and H. Meng. 1975. Falcons return. 
Wilham Morrow and Co., New York. 

Mebs, T. 1969. Wanderfalkenbruten an menschlichen 
Bauwerken. Deutscher Falkenorden 1968: 55-65. 

Mebs, T. 1988. The return of the Peregrine Falcon in 
West Germany. Pages 173-178 in Peregrine Falcon 
populations: their management and recovery. Edited 
by T. J. Cade, J. H. Enderson, C. G. Thelander and 
C. M. White. The Peregrine Fund, Inc., Boise, Idaho. 

Peakall, D. B., D. G. Noble, J. E. Elliott, J. D. Somers, 
and G. Erickson. 1990. Environment contaminants 
in Canadian Peregrine Falcons, Falco peregrinus: a 
toxicological assessment. Canadian Field-Naturalist 
104(2): 244-254. 

Penny, L. 1981. Peregrine fever. The East Hampton 
Star, 3 September. 

Saar, C. 1988. Reintroduction of the Peregrine Falcon 
in Germany. Pages 629 636 in Peregrine Falcon 
populations: their management and recovery. Edited 
by T. J. Cade, J. H. Enderson, C. G. Thelander and 
C. M. White. The Peregrine Fund, Inc., Boise, Idaho. 

Tansy, M. F., and R. P. Roth. 1970. Pigeons: a new 
role in air pollution. Journal of the Air Pollution 
Control Association 20: 307-309. 



Received 8 June 1989 
Accepted 9 May 1990 



Impact of Forced Renesting on Reproductive Success in 
Ungava Bay Peregrine Falcons, Falco peregrinus 



David M. Bird', Ian Ritchiei, James D. Weaver2, and Reed Bowman^ 

'Macdonald Raptor Research Centre of McGill University, 21,111 Lakeshore Road, Ste. Anne de Bellevue, Quebec 

H9X ICO 
2The Peregrine Fund, Inc., 5666 West Flying Hawk Lane, Boise, Idaho 83709 
^National Audubon Society, Research Department, 115 Indian Mound Trail, Tavernier, Florida 33070 

Bird, David M., Ian Ritchie, James D. Weaver, and Reed Bowman. 1990. Impact of forced renesting on reproductive 
success in Ungava Bay Peregrine Falcons, Falco peregrinus. Canadian Field-Naturalist 104(2): 219-221. 

The first clutches were removed from eight pairs of the Peregrine Falcon nesting in Ungava Bay in 1984 and 1986 to 
induce the laying of a replacement clutch, but only one pair renested to produce two young. 

En 1 984 puis en 1 986, la premiere couvee de huit couples de Faucons pelerins qui nidifiaient dans la Bale d'Ungave a ete 
enlevee dans le but d'inciter la ponte d'une couvee de remplacement. Un couple seulement a renidifie et a produit deux 
jeunes. 

Key Words: Peregrine Falcon, Falco peregrinus, forced renesting, replacement clutch, Ungava Bay. 



Many raptors can produce a second or 
replacement clutch upon loss of their first 
(Olendorff 1971; Morrison and Walton 1980). The 
technique of removing the first clutch to induce the 
laying of a second or even a third clutch was 
discovered by egg-collectors many years ago (Bent 
1938). 

Forced renesting, the technique of removing the 
first clutch to induce the laying of a second or even 
a third clutch, has been shown to have little impact 
on egg dimensions, fertility or hatchabiUty in wild 
American Kestrels {Falco sparverius) (Bowman 
and Bird 1985). However, in captive kestrels 
seasonal declines were noted for fertility, 
hatchability, clutch size, egg length, eggshell 
thickness, and fresh egg weight (Bird and Lague 
1982). 

Forced renesting is commonly used to enhance 
reproductive output in captive breeding programs 
for many species of birds of prey, most notably the 
endangered Peregrine Falcon {F. peregrinus) 
(Weaver and Cade 1983). Moreover, it has 
occasionally been used successfully as a manage- 
ment tool for wild peregrine nesting at southerly 
latitudes (Barclay 1988). While Peregrine Falcons 
are making a strong comeback in the eastern 
United States due mainly to an intensive captive 
breeding and release program (Barclay 1988), the 
number of successfully breeding pairs continues to 
remain low in eastern Canada despite similar 
programs (Holroyd and Banasch, this issue). 

Ungava Bay in northern Quebec has supported a 
stable population of Peregrine Falcons from 1980 
to 1983 (Bird and Weaver 1988). Forcing Ungava 
peregrines to lay replacement clutches, while 
artificially raising the progeny from first clutches, 



would provide an inexpensive source of falcon 
stock for release programs in southern Quebec, 
given that replacement clutches are as successful as 
first clutches. 

With this in mind, we collected first clutches 
from three pairs of peregrines in 1984 and five pairs 
in 1986 from the Koksoak River north of 
Kuujjuaq, Quebec (latitude 58° 15') and Leaf Bay 
(latitude 58° 45'). Nests were visited by helicopter 
on 9 June 1984 and 15 June 1986, respectively, and 
the eggs kept warm in a vinyl suitcase lined with 
soft foam and a hot water bottle. The eggs were 
subsequently transferred to a Marsh Farms Roll-X 
incubator (College Pets and Poultry Supplies, 
Toronto) and kept at a standard temperature and 
humidity regime for peregrine eggs (Weaver and 
Cade 1983). After a day or two, the eggs were again 
placed in the vinyl case with hot water bottle and 
flown by commercial airline south to the 
Macdonald Raptor Research Centre where they 
were transferred to Marsh Farms Roll-X 
incubators until hatching. All young in both years 
were fed four times daily on frozen-thawed, day- 
old cockerels and later released at sites in southern 
Quebec. 

On 20 July 1984 and on 23 August 1986, 
respectively, we returned to the sites where clutches 
were removed to determine whether renesting 
occurred. We also examined possible alternate 
sites in the surrounding area. With one exception, 
the falcons were either absent from the area or they 
exhibited little no defensive behaviour, indicating 
a failure to renest (Table 1). The pair that did renest 
strongly defended two healthy young. Assuming 
an incubation period of 32 days and a laying 
interval of 2 days between eggs, the nestlings' age of 



219 



220 



The Canadian Field-Naturalist 



Vol. 104 



Table 1. Impact of removal of first clutches on subsequent reproductive success of Peregrine Falcons nesting in 
Ungava Bay in 1984 and 1986. 











No. days 




No. 


No. 


No. 


No. of 






Initiation 


Date of 


natural 


No. 


eggs 


eggs 


young 


young in 


Year 


Site 


of clutch* 


removal 


incubation* 


eggs 


fertile 


hatched 


fledged 


2nd clutch 


1984 


A 


30 May 


6 June 


6 


3 


3 


1 


1 


— 


1984 


B 


28 May 


1 1 June 


10 


3 


3 


2 


2 


— 


1984 


C 


2 June 


1 1 June 


3 


4 


1 








— 


1986 


D 


24 May 


17 June 


18 


4 


4 


4 


4 


— 


1986 


E 


28 May 


17 June 


14 


4 


4 


4 


4 


— 


1986 


F 


2 June 


17 June 


11 


3 


1** 


1 





— 


1986 


G 


5 June 


15 June 


5 


4 


4 


3 


3 


2 


1986 


H 


7 June 


17 June 


6 

Totals 


3 


3 


3 


3 


— 




28 


23 


18 


17 


2 



* All but C are approximations based on the known hatching date and assuming an incubation period of 32 days. The 
data for C were calculated on measured egg weight loss based on Burnham (1983). 
**The other two eggs were addled and fertility could not be determined. 



approximately 3 weeks indicates a renesting 
interval of roughly 2'/2 to 3 weeks. This concurs 
with the findings of Ratcliffe ( 1 980) who calculated 
a mean renesting interval of 19 to 20 days for 43 
renesting pairs of Peregrine Falcons in Great 
Britain. 

Ungava peregrines were consistent in their 
timing for initiating laying between the two years 
(Table 1). Not consistent however, was the number 
of days from the date of the last egg laid to the 
removal of the clutch. Peregrines are most likely to 
renest if the first clutch is lost before 7 to 10 days of 
incubation (Ratcliffe 1980). After 10 days, the 
external stimuli, e.g. presence of a mate, suitable 
nesting site, food, etc. apparently are not strong 
enough to create the physiological conditions 
necessary for egg production and laying. Although 
some pairs on rare occasions have laid second 
clutches after incubating the first one full-term 
(Ratcliffe 1980), this would appear to be highly 
unlikely for northern-nesting falcons. 

Successful laying of replacement clutches that 
fledged young has been documented for Gyrfal- 
cons {F. rusticolus) at latitudes between 60 and 69 
degrees (Piatt 1976; Poole 1988). However, based 
on anecdotal observations by Hickey (1942) and 
Cade (I960), renesting attempts by peregrines at 
high latitudes have generally not been successful. 
The less migratory Gyrfalcons are often 2 to 3 
weeks ahead of peregrines in their breeding 
chronology in Ungava Bay (D. Bird, unpublished 
data). The highly migratory northern peregrine 
populations have a much shorter breeding period 
in which to achieve pair-integration prior to 
breeding compared to sedentary populations 
(Fischer 1967). Also, whether the nestling period 
for second clutches exceeds that of first clutches in 
peregrines is not known. Bowrnan and Bird (I9H5) 
noted that second clutch young of wild kestrels 
took longer to fledge than first clutch ones. 



Certainly young peregrines fledging from 
northern nests would require enough time to hone 
flight and hunting skills before migrating to parts as 
far south as Argentina. It is questionable whether 
the two young produced by the renesting falcons 
could achieve that. Being three weeks of age on 24 
August, they would still be practising foraging skills 
in Ungava Bay well into late September. 

Egg hatchability was 78 percent (18/23), 
indicating little, if any, effect of handling them 
throughout the study on incubation success. 
Considering only those eggs receiving anywhere 
from 6 to 18 days of natural incubation, the 
hatchability rose to 100 percent. Lowered 
hatchability in artificially-incubated eggs of 
captive peregrines without any prior natural 
incubation has been well-documented (Burnham 
1983). The only way to assure a period of 7 to 10 
days natural incubation in northern peregrines 
would be to actually observe the nesting pair to 
determine egg-laying dates beforehand. This is 
somewhat impractical for most nest sites in 
Ungava Bay due to hazardous, unpredictable 
weather conditions, not to mention the cost factor. 

Our findings concur with earlier observations by 
Hickey (1942) and Cade ( 1 960). We conclude then 
that removal of first clutches to induce the laying of 
a second one is not a practical method of 
augmenting the numbers of Peregrine Falcons to 
be released in southern Canada. 

Acknowledgments 

Support for this study was generously provided 
by the World Wildlife Fund (Canada), the 
Ministry of Recreation, Fish and Game of Quebec, 
The Peregrine Fund, Inc., and Canadian Airlines. 
We are also very grateful to J. Grist, S. and E. 
Gordon, W. Cooper, A. Kudluk, T. Hrictic, as well 
as the towns of Kuujjuaq and Tasiuq for their kind 
help. 



1990 



Bird, Ritchie, Weaver, and Bowman: Forced Renesting 



221 



Literature Cited 

Barclay, J. H. 1988. Peregrine restoration in the eastern 
United States. Pages 549-558 in Peregrine Falcon 
populations, their management and recovery. Edited by 
T. J. Cade, J. H. Enderson, C. G. Thelander, and C. M. 
White. The Peregrine Fund, Inc., Boise, Idaho. 

Bent, A. C. 1938. Life histories of North American birds 
of prey. Part 2. Orders Falconiformes and Strigiformes. 
Dover Publications, New York. 482 pages. 

Bird, D. M., and P. C. Lague. 1982. Influence of forced 
renesting, seasonal date of laying, and female character- 
istics on clutch size and egg traits in captive American 
Kestrels. Canadian Journal of Zoology 60: 71-79. 

Bird, D. M., and J. D. Weaver. 1988. Peregrine Falcon 
populations in Ungava Bay, Quebec, 1980-1985. In 
Peregrine Falcon populations, their management and 
recovery. Edited by T. J. Cade, J. H. Enderson, C. G. 
Thelander, and C. M. White. The Peregrine Fund, Inc., 
Boise, Idaho. 

Bowman, R., and D. M. Bird. 1985. Reproductive 
performance of American Kestrels laying replacement 
clutches. Canadian Journal of Zoology 63: 2590-2593. 

Burnham, W. A. 1983. Artificial incubation of falcon 
eggs. Journal of Wildlife Management 47: 158-168. 

Cade, T. J. 1960. Ecology of the peregrine and Gyrfalcon 
populations in Alaska. University of California 
Publications in Zoology 63: 151-290. 



Fischer, W. 1967. Der Wanderfalk. A. Ziemsen Verlag, 

Wittenberg Lutherstadt. 262 pages. 
Hickey, J.J. 1942. Eastern populations of the Duck 

Hawk. Auk 59: 176-204. 
Holroyd, G. L. and U. Banasch. 1990. The reintroduc- 

tion of the Peregrine Falcon, Falcoperegrinus, anatum 

into southern Canada. Canadian Field-Naturalist 

104(2): 203-208. 
Morrison, M. L. and B. J. Walton. 1980. The laying of 

replacement clutches by falconiforms and strigiforms 

in North America. Raptor Research 14: 79-85. 
Olendorff, R. R. 1971. Falconiform reproduction; a 

review. Part 1. Raptor Research Report No. 1. 

Vermillion, South Dakota. 1 1 1 pages. 
Piatt, J. B. 1976. Gyrfalcon nest site selection and 

winter activity in the western Canadian arctic. 

Canadian Field-NaturaUst 90: 338-345. 
Poole, K. G. 1988. A replacement clutch in wild 

Gyrfalcons, Falco rusticolus, in the Northwest 

Territories. Canadian Field-NaturaHst 102: 62-64. 
Ratcliffe, D. 1980. The Peregrine Falcon. Buteo Books, 

Vermillion, South Dakota. 416 pages. 
Weaver, J. D. and T.J. Cade. Editors. 1983. Falcon 

propagation: a manual on captive breeding. The 

Peregrine Fund, Inc., Ithaca, New York. 100 pages. 

Received 8 June 1989 
Accepted 9 May 1990 



Levels of Contaminants in Canadian Raptors, 1966 to 1988; Effects 
and Temporal Trends 

David G. Noble and John E. Elliott 

Canadian Wildlife Service, Ottawa, Ontario KIA 0H3 

Noble, David G., and John E. Elliott. 1990. Levels of contaminants in Canadian raptors, 1966 to 1988; effects and 
temporal trends. Canadian Field-Naturalist 104(2): 222-243. 

Organochlorine and mercury residue concentrations in eggs and body tissues of twenty-seven species of raptors 
collected in Canada between 1966 and 1988 are summarized. A few individuals had liver mercury or DDE 
concentrations at levels associated with poisoning. During the late 1960s and early 1970s, levels of DDE in some eggs 
of Bald Eagles, Osprey, Red-tailed Hawks, Northern Harriers, Merlins, Prairie Falcons, and Great Horned Owls were 
high enough to cause eggshell thinning. In the 1980s, elevated DDE levels were found in some eggs of Sharp-shinned 
Hawks, Cooper's Hawks and Merlins. Dieldrin and heptachlor epoxide levels in eggs also exceeded minimum critical 
levels in a few individuals of a number of species. The significance of the detected concentrations are discussed in 
relation to the status of those populations. DDE, dieldrin and heptachlor epoxide concentrations in eggs generally 
declined over the period of sampling. Although some bird-eating raptors are still highly contaminated, population 
declines in many species can be attributed to a number of factors, including loss of suitable habitat. 

Les concentrations de residus de mercure et d'organochlores dans les oeufs et les tissus corporels de vingt sept 
especes de rapaces ramassees au Canada entre 1966 et 1988 sont compiles. Quelques individus avaient des 
concentrations de mercure ou de DDE dans le foie a des niveaux associes aux empoisonnements. Vers la fin des annees 
60 et au debut des annees 70, les niveaux de DDE dans certains oeufs de Pygargue a tete blanche, de Balbuzard, de Buse 
a queue rousse, de Busard Saint-Martin, de Faucon emerillons, de Faucon des Prairies et de Grand-due d'Amerique 
etaient assez eleves pour causer un amincissement de la coquille. Pendant les annees 80, des taux de DDE eleves ont ete 
trouves dans certains oeufs d'Epervier brun, d'Epervier de Cooper et de Faucon emerillon. Les niveaux de dieldrine et 
d'epoxyde d'heptachlore dans les oeufs depassaient aussi les taux critiques minimums dans quelques specimens d'un 
certain nombre d'especes. L'importance des concentrations relevees est discutee en fonction du statut des populations 
en question. Les concentrations de DDE, de dieldrine et d'epoxyude d'heptachlore dans les oeufs diminuaient, en 
general, pendant la periode d'echantillonage. Meme si les rapaces mangeurs d'oiseaux sont encore considerablement 
contamines, le declin des populations de beaucoup d'especes peut etre attribue a un certain nombre de facteurs dont la 
perte d'un habitat adequat. 

Key Words: Raptors, Canada, organoclorines, mercury, DDE concentrations, diedrin, heptachlor epoxide, 
population declines. 

Since the discovery of synthetic pesticides such Falcon, Falco peregrinus, particularly the anatum 

as DDT and dieldrin in wildlife, birds of prey have race, which had been extirpated from nearly all of 

been identified as being particularly vulnerable its former breeding range in eastern North 

(Newton 1979). Because of their position at the top America (Hickey 1969). 

of terrestrial or aquatic food chains, and their Use of the high-molecular-weight organochlo- 

relatively inefficient elimination of lipophilic rine pesticides (such as DDT, dieldrin, heptachlor, 

contaminants (Walker and Stanley 1986), raptors chlordane, BHC and mirex) and many applica- 

tend to accumulate high levels of those substances. tions of mercury have been restricted in Canada 

Moreover, many species appear to be relatively and the United States since the early 1970s, 

sensitive to the eggshell-thinning effects of DDE However, there continue to be reports of high 

(Cooke, 1973; Fyfe et al. 1988) and to the direct levels in wildlife (e.g., Clark and Krynitsky 1983; 

toxic effects of other organochlorines such as Blus et al. 1987). This paper summarizes data on 

dieldrin (Lockie et al. 1969). environmental contaminants in raptors collected 

Early indications of the dangers posed by DDT by the Canadian Wildlife Service (CWS) since 

and other organochlorines to North American 1966. A more detailed breakdown of the data is 

raptors included the rapid declines in numbers of available in a CWS Technical Report (Noble and 

Osprey Pandion haliaetus in the eastern United EWiolt in press). 
States (Ames 1966) and Bald Eagles, Haliaeetus 

leucocephalus in Ontario (Postupalsky 1971), in Methods 

conjunction with reports of thin-5;hclled eggs that The data were retrieved from the CWS National 

either disappeared or failed to hatch, in 1965, Registry of Toxic Chemical Residues (Elliott et al. 

attention focussed on the plight of the Peregrine 1987), a computerized repository of data on 

222 



1990 Noble and Elliot: Levels of Contaminants in Canadian Raptors 



223 



Table 1 . Summary of minimum critical levels (in mg/ kg, wet weight) of major environmental contaminants in eggs, 
brains and livers of raptors. 



Conntaminant Level (mg/kg) diagnostic of: 



(1) acute toxicity 
Brain Liver 



(2) reproductive effects 
Eggs 



Dieldrin 

Oxychlordane 

Heptachlor epoxide 

DDE 

PCBs 

HCB 

Mercury 



>5.0^ 


3-1 Ob 


> 1.0^ 


1.1-5.0" 


3-10" 




3.4-8.31 


3-10" 


>1.5^ 


250f 


100" 


1.2-308 


500-3000" 




>50' 
>5.0J 


>50'' 


20-45'= 


>0.5' 



^Stickel et al. 1969;''Cookeet al. 1982;^Lockieetal. 1969; Wiemeyeret al. 1975;dStickelet al. 1979;<=Henny et al. 1983; 
fStickel et al. 1984; sFyfe et al. 1988, Lincer 1975; hReath et al. 1972; iHoffman et al. 1987; JBoersma et al. 1986; 
kScheuhammer 1987; 'Fimreite 1971; Heinz 1979. 



environmental contaminants in Canadian wildlife. 
We obtained data on nineteen species of hawks, 
eagles and vultures and eight species of owls. The 
contaminants selected include DDE (the main 
metabolite of DDT), dieldrin, heptachlor epoxide 
(the major metabolite of heptachlor), oxychlor- 
dane (a metabolite of cis and trans chlordane), 
hexachlorobenzene (HCB), polychlorinated 
biphenyls (PCBs) and mercury. 

Data are presented on the basis of wet weight. 
This minimizes variation due to changes in lipid 
content during development of eggs (Peakall and 
Oilman 1979) or due to physiological factors 
affecting levels in tissues. Concentrations in 
dehydrated eggs were adjusted for estimated water 
loss. Results which were below the detection limit 
were assigned a value of one half the detection 
limit, normally 0.005 mg/kg (O.E.CD. 1981). 
Geometric means were determined for samples 
grouped by time period of collection and 
geographic region. The three time periods 
represent, respectively, the era of extensive 
organochlorine use ( 1 965- 1 972), immediately post- 
ban (1973-1979) and the recent situation (1980- 
1988). 

For species for which sufficient data were 
available, we investigated changes in contaminant 
levels over time by conducting pair-wise t-tests of 
samples grouped by region. In addition, we used 
the general hnear model (GLM) procedure of S AS 
to test for a significant directional trend. 

Methods of collection of the specimens varied 
over the twenty-three year period. Many eggs were 
collected opportunistically at the end of the 
breeding season, and therefore represent mainly 
nonviable eggs. However, many fresh eggs, 
particularly those of Merlins, Falco columbarius 
and Prairie Falcons, Falco mexicanus, were also 
collected. Unless stated otherwise, eggs analyzed 
were from separate nests. 

Tissues were usually obtained from birds found 
dead. Although breast muscle, fat, gonads and 



whole bodies were also analyzed, only liver and 
brain residues are reported here. Further details of 
the sample collection, preparation and storage are 
described in Noble and Elliott {in press). 

The earliest data included are from 1966. Prior 
to 1969, only DDE and dieldrin concentrations are 
presented, due to the confounding influence of 
PCBs at that time. A method of separating peaks 
of PCBs from those of other organochlorines was 
developed on behalf of CWS in 1969 (Reynolds 
1969). We have presented the PCB data on the 
basis of a 1:1 ratio of Aroclors 1254 and 1260. 
Total mercury was determined by wet digestion 
followed by flameless atomic absorption spectro- 
photometry (AAS), as described by Hatch and Ott 
(1968). 

Until 1984, all analyses were carried out at the 
Ontario Research Foundation, following proce- 
dures described by Reynolds and Cooper (1975). 
Since 1984, samples have been analyzed at the 
National Wildlife Research Centre in Hull, using 
methods described in Peakall et al. (1986). 

Results and Discussion 

Evaluation of the effects of detected contaminants 
The objective of this section is to compare the 
detected concentrations with critical values derived 
from the literature. Minimum critical levels and 
their sources are summarized in Table 1, and 
discussed in more detail in Peakall et al. {this issue). 
The acute lethal toxicities of organochlorines 
and mercury have been experimentally determined 
in few raptor species: Goshawks, Accipiter gentilis 
(Borg et al. 1970), Red-tailed Hawks, Buteo 
jamaicensis (Fimreite and Karstad 1971) and 
American Kestrels, Falco sparverius (Porter and 
Wiemeyer 1972). Hence, we supplemented the 
findings of those studies with experimental work 
on other birds (Stickel et al. 1969; Heath et al. 
1972; Stickel et aL 1979; Cooke et al. 1982; Stickel 
et al. 1984), to derive the critical brain and liver 
values in Table 1. 



224 



The Canadian Field-Naturalist 



Vol. 104 



Laboratory studies on the effects of organochlo- 
rines on reproduction in raptors have been limited 
to work on Screech Owls, Otus asio (McLane and 
Hughes 1980), Barn Owls, Tyto alba (Mendenhall 
et al. 1983) and American Kestrels (e.g., Wiemeyer 
et al. 1986). Minimum critical values were 
determined from the results of those and other 
studies on the relationship between contaminant 
levels in eggs and reproductive parameters (e.g., 
Lockie et al. 1969; Enderson and Berger 1970; 
Snyder et al. 1973; Wiemeyer et al. 1975; Henny et 
al. 1983; Hoffman et al. 1987; Fyfe et al. 1988). 
Note that because of possible covariance between 
contaminants or with environmental effects, a 
cause and effect relationship cannot be assumed in 
most field studies. 

Incidences of raptor mortality 
due to contaminants 

Concentrations in brains and livers are 
presented in Appendix lA, IB. A very small 
proportion of the raptor tissue samples analyzed 
were found to contain any contaminants at levels 
diagnostic of acute toxicity. In 1970, the liver of a 
Bald Eagle from northern Ontario was found to 
contain 42 mg/ kg mercury, and the liver of one of 
three Turkey Vultures, Cathartes aura, collected in 
the same year contained 60 mg/ kg mercury. In the 
early 1980s, DDE in the hver of one of seven 
Sharp-shinned Hawks, Accipiter striatus, col- 
lected in British Columbia exceeded 100 mg/kg, 
the critical value proposed by Cooke et al. (1982). 
Although not diagnostic of pesticide poisoning, 
the brains of two American Kestrels found dead in 
Ontario in 1987 contained dieldrin in excess of 
2 mg/kg and heptachlor epoxide in excess of 
1 mg/kg (Barker, personal communication). 

As there has traditionally been no formal system 
in place to retrieve and analyze raptors found dead, 
we cannot accurately assess the incidence of raptor 
mortality due to contaminants. Although the 
infrastructure for testing dead wildlife for toxic 
substances has improved, most poisoned birds are 
seldom recovered. Hence, actual mortality rates 
are undoubtedly higher than those observed. 

Poisoned raptors have been reported more 
frequently in the United States than in Canada. 
Dieldin was implicated in the deaths of Bald Eagles 
(Kaiser et al. 1980), Peregrine Falcons (Reichel et 
al. 1974) and Red-shouldered Hawks, Buteo 
lineatus (Sundlof et al. 1986). Oxychlordane was 
considered to be the cause of the deaths of two 
Red-shouldered Hawks and a Great Horned Owl, 
Buho virf^inianus (Blus et al. 1983) and heptachlor 
epoxide was implicated in the deaths of three 
Golden Eagles, Aquila chrysaetos, an American 
Kestrel, and a Rough-legged Hawk, Buteo lagopus 
(Henny et al. 1984). In 1980, a Cooper's Hawk, 
Accipiter cooperi, was found dead in Maryland 



with lethal concentrations of DDT in its brain 
(Prouty et al. 1982). In New York, two of twenty 
Great Horned Owls examined were considered to 
have been poisoned by either DDE, dieldrin or 
PCBs (Stone and Okoniewski 1983), and an 
American Kestrel was found to contain toxic 
concentrations of both DDT and dieldrin in its 
brain (Stone 1981). Many of these deaths may 
involve Canadian birds. A Merlin found dead in 
New Mexico with more than 9 mg/kg heptachlor 
epoxide in its brain was found to be banded in 
Alberta (Henny et al. 1976). 

Although not documented in North America, 
poisoning by mercury has been reported in 
European raptors (Borg et al. 1969; Delbeke et al. 
1984). 

Levels and reproductive effects 

During the late 1960s and early 1970s, a number 
of Canadian raptor populations were found to 
have elevated levels of organochlorines in their 
eggs. By the 1980s, significant contamination was 
reported only in eggs of Sharp-shinned Hawks, 
Cooper's Hawks, Merlins and Peregrine Falcons. 

The pattern of contamination by species {see 
Figure 1 for recent data) confirms the maxim that 
species which feed primarily on migratory birds are 
most contaminated, whereas species feeding 
mainly on mammals are least contaminated. Local 
pesticide use, food from aquatic sources and 
wintering range may also be important factors 
(Noble and Elliott in press). 

Accipiters were relatively contaminated. 
Between 1980 and 1988, three of thirteen Cooper's 
Hawk eggs and five of nine Sharp-shinned Hawk 
eggs collected in southern Ontario contained DDE 
in excess of 10 mg/ kg (Table 2). DDE egg residues 
of this magnitude have been associated with 
significant eggshell thinning in North American 
accipiters (Snyder et al. 1973) and Eurasian 
Sparrowhawks Accipiter nisus (Newton et al. 
1986). During the same time period, dieldrin was 
detected at a level greater than 1 mg/kg in one 
Sharp-shinned Hawk egg. Throughout this paper, 
we have used 1 mg/kg dieldrin as the minimum 
critical level in the eggs of raptors, based on the 
studies of Lockie et al. (1969) and Wiemeyer et al. 
(1975). However, there may be considerable 
interspecific differences in susceptibility, as 3 mg/ 
kg dieldrin was found to have no effect on 
reproduction in Screech Owls (Mendenhall et al. 
1983). 

Snyder ct al. (1973) demonstrated a significant 
correlation between the percentage of birds in the 
diet of individual Cooper's Hawks and the DDE 
content of their eggs. Of the accipiters. Sharp- 
shinned Hawks depend most on migratory 
insectivorous birds, whereas Northern Goshawks 
feed mainly on resident birds and mammals 



1990 Noble and Elliot: Levels of Contaminants in Canadian Raptors 



225 



Bald Eagle 




2.311 (2) 








Sharp-shinned Hawk 




8.2848 


(12) 














Cooper's Hawk 




4.613 (13) 






N. Goshawk 




0.991 (4) 








Amer. Kestrel 




3.665 (5 pools) 




Great Grey Owl 


0.059 (3) 









2 4 6 
DDE (mg/kg) 


8 10 


Merlin 




2.953 (14) 




Prairie Falcon 
Gyrfalcon 

Rough-legged Hawk 
Swainson's Hawk 
Ferruginous Hawk 


0. 
0. 


0.384 (16) 
124 (6) 
0.329 (6) 
0.397 (8 pools) 
02 (2 pools) 









0.0 0.5 1.0 1.5 2.0 2.5 3.0 3.5 4.0 

DDE (mg/kg) 

Figure 1. Comparison of DDE residue levels in eggs of raptors during the 1980s; A: 
from Ontario and east during the 1960s; B: from western Canada. Values are 
geometric means and sample sizes, the latter in parenthesis). 



(Sherrod 1978). Comparison of the organochlo- 
rine content of eggs of all three accipiters (see 
Appendix 2A) is consistent with that hypothesis. 
Overall, buteos were considerably less contami- 
nated than accipiters (Appendix 2B). Four Red- 
tailed Hawk eggs collected in 1968 were the only 
eggs to contain DDE residues in excess of 15 mg/ 
kg (Table 2), a level associated with minor eggshell 
thinning in this species (Seidensticker and 
Reynolds 1971). Dieldrin concentrations exceeded 
1 mg/kg in five Red-tailed Hawk eggs, two 
Swainson's Hawk, Buteo swainsoni, eggs and two 
Ferruginous Hawk, Buteo regalis, eggs, all 
collected in the late 1960s (Table 2). Heptachlor 
epoxide levels exceeded the minimum critical value 
of 1.5 mg/kg(Henny et al. 1983) in one Red-tailed 
Hawk egg, one Swainson's Hawk egg, one 
Ferruginous Hawk egg and one Rough-legged 



Hawk egg (Table 2). However, there is some 
evidence that Swainson's Hawks, at least, can 
tolerate concentrations in excess of 2.5 mg/kg 
(Henny et al. 1984). One Red-tailed Hawk egg also 
contained 1.6 mg/kg mercury, slightly more than 
the level associated with detrimental reproductive 
effects in other birds (Fimreite, 1971; Heinz 1979). 
Low levels of DDE in eggs of Ferruginous and 
Swainson's hawks in the Dakotas, during the 
1970s, were attributed to a diet of small mammals 
(Stendell et al. 1988). Small mammmals dominate 
the diet of most buteos (Lincer et al. 1971; Schmutz 
and Schmutz, 1980; Risley 1983). As in Stendell et 
al.'s (1988) and Bechard's (1981) study, contami- 
nant residues in Swainson's Hawks were similar to 
those in coexisting Ferruginous Hawks, despite the 
fact that the former species is highly migratory, 
wintering south to Argentina (Palmer 1988). 



226 



The Canadian Field-Naturalist 



Vol. 104 



Table 2. Proportion of eggs with threshold critical levels of DDE, dieldrin, heptachlor expoxide or mercury, likely to 
affect productivity. The critical values are 10 mg/kg DDE (except where indicated), 1 mg/kg dieldrin, 1.5 mg/kg 
heptachlor expoxide and 0.5 mg/kg total mercury. No pooled samples are included. 



Region 


Years 


DDE 


Dieldrin 


HE 


Mercury 


Northern Goshawk 












Atlantic 


1965-72 


0/1 


0/1 


0/1 


- 


Ontario 


1980-88 


0/4 


0/4 


0/4 


- 


Cooper's Hawk 












Ontario 


1980-88 


3/13 


0/13 


0/13 


- 


Prairies 


1965-72 


1/17 


1/17 


0/17 


0/7 




1973-79 


0/3 


1/3 


0/3 


0/1 


Sharp-shinned Hawk 












Atlantic 


1980-88 


0/3 


0/3 


0/3 


- 


Ontario 


1980-88 


5/9 


1/9 


0/9 


- 


Prairies 


1965-72 


0/1 


0/1 


0/1 


0/1 


Red-tailed Hawk 












Ontario 


1965-72 


0/1 


0/1 


0/2 


0/2 


Prairies 


1965-72 


4/46 


5/46 


1/42 


1/14 


British Columbia 


1965-72 


0/1 


0/1 


0/1 


- 


Red-shouldered Hawk 












Ontario 


1973-79 


0/3 


0/3 


0/3 


- 


Swainson's Hawk 












Prairies 


1965-72 


0/18 


2/17 


1/17 


- 




1973-79 


0/1 


0/1 


0/1 


0/1 


Ferruginous Hawk 












Prairies 


1965-72 


1/50 


2/49 


1/50 


- 


Rough-legged Hawk 












North 


1965-72 


0/13 


0/13 


0/3 


0/3 




1973-79 


0/1 


0/1 


0/1 


- 




1980-88 


0/6 


0/6 


1/6 


0/2 


Bald Eagle' 












Atlantic 


1965-72 


1/1 


0/1 


- 


0/1 




1980-88 


0/2 




- 




Ontario 


1965-72 


16/16 


10/16 


0/14 


5/15 




1973-79 


2/2 


1/2 


0/2 


0/2 


Prairies 


1965-72 


4/13 


2/13 


0/5 


1/10 


Golden Eagle 












Prairies 


1965-72 


0/9 


0/9 


0/9 


0/6 




1973-79 


0/12 


1/12 


0/12 


0/12 


North 


1973-79 


0/1 


0/1- 


- 


- 


Osprey2 












Atlantic 


1965-72 


3/3 


0/1 


- 


0/3 


Ontario 


1965-72 


4/8 


0/8 


0/1 


0/8 


Prairies 


1973-79 


1/2 


0/2 


0/2 


0/2 


British Columbia 


1965-72 


1/2 


0/2 


0/2 


- 


North 


1973-79 


2/2 


0/2 


0/2 


0/2 


Northern Harrier 












Quebec 


1965-72 


1/1 


0/1 


0/1 


0/1 


Prairies 


1965-72 


3/22 


3/22 


1/22 


0/8 


Merlin' 












N. Quebec 


1965-72 


l/I 


0/1 


0/1 


l/I 


Prairies 


1965-72 


88/165 


32/165 


13/165 


8/93 




1973-79 


129/159 


19/159 


23/159 


1/141 




1980-88 


5/14 


0/14 


3/14 


- 


British Columbia 


1965-72 


1/1 


0/1 


0/1 


3/3 


North 


1973-79 


4/4 


2/4 


0/4 


0/4 


Prairie Falcon'' 












Prairies 


1965-72 


154/249 


17/249 


9/249 


17/229 




1973-79 


132/216 


12/216 


16/216 


0/180 




1980-88 


3/16 


6/16 


0/16 


0/12 


Peregrine Falcon ianaium) 












Atlantic 


1965-72 


1/1 


0/1 


0/1 


0/1 


Quebec 


1980-88 


4/7 


3/7 


0/7 


- 



1990 



Noble and Elliot: Levels of Contaminants in Canadian Raptors 227 



Table 2. Concluded 



Region 


Years 


DDE 


Dieldrin 


HE 


Mercury 


Prairies 


1965-72 


5/5 


3/5 


0/5 


1/5 




1973-79 


12/20 


2/20 


0/20 


1/5 




1980-88 


8/14 


3/14 


3/14 


3/11 


North 


1965-72 


6/6 


1/6 


0/6 


0/6 




1973-79 


4/4 


2/4 


0/4 


1/3 




1980-88 


6/9 


1/9 


1/9 


- 


Peregrine Falcon {tundrius) 












Quebec 


1965-72 


4/10 


4/10 


- 


- 




1973-79 


3/4 


3/4 


0/4 


1/4 




1980-88 


2/10 


2/10 


0/10 


- 


North 


1965-72 


7/23 


8/22 


0/15 


5/23 




1973-79 


13/20 


4/20 


3/20 


0/19 




1980-88 


5/19 


4/19 


3/19 


0/8 


Peregrine Falcon (pealei) 












British Columbia 


1965-72 


7/12 


0/12 


0/6 


7/10 




1980-88 


0/4 


0/4 


0/4 


- 


Gyrfalcon 












North 


1965-72 


0/1 


0/1 


0/1 


1/1 




1973-79 


0/10 


0/10 


0/10 


0/4 




1980-88 


0/6 


1/6 


0/6 


1/1 


American Kestrel 












Atlantic 


1965-72 


0/1 


0/1 


- 


- 


Prairies 


1965-72 


1/8 


1/8 


1/8 


0/3 


Great Horned Owl 












Ontario 


1965-72 


0/1 


0/1 


- 


- 


Prairies 


1965-72 


6/30 


0/30 


2/29 


0/7 


North 


1965-72 


0/1 


0/1 


0/1 


1/1 


Snowy Owl 












North 


1973-79 


0/1 


0/1 


0/1 


- 


Burrowing Owl 












Prairies 


1965-72 


0/1 


0/1 


0/1 


0/1 


Great Grey Owl 












Minnesota 


1980-88 


0/3 


0/3 


0/3 


- 


Long-earned Owl 












Prairies 


1965-72 


0/7 


0/7 


0/7 


0/3 


Short-eared Owl 












Prairies 


1965-72 


0/6 


0/6 


0/6 


- 



'Bald Eagle: critical value of DDE = 6 mg/kg; ^Osprey: critical value of DDE 
DDE = 5 mg/kg; "Prairie Falcon: critical value of DDE = 1.2 mg/kg. 



4 mg/kg; ^Merlin: critical value of 



Bald Eagles, which feed on a variety offish, birds 
and carrion, had high concentrations of contami- 
nants in their eggs (Appendix 2C). Twenty-three of 
34 eggs contained DDE residues in excess of 6 mg/ 
kg (Table 2), a value associated with significant 
eggshell thinning in this species (Wiemeyer et al. 
1984). Maximum values of DDE were 100 mg/ kg, 
in areas of northern Ontario where reproductive 
failures were common (Grier 1974). Six eggs 
contained more than 2 mg/ kg dieldrin. Concentra- 
tions of heptachlor epoxide and other organochlo- 
rines were relatively low, and mercury exceeded 1 
mg/kg in only two eggs (Table 2). 

In eggs of Golden Eagles, which feed primarily 
on mammals (Sherrod 1978), only dieldrin (at 
2.12 mg/kg in one egg from Alberta) was found at 
concentrations associated with reduced productiv- 



ity in a Scottish eagle population (Lockie et al. 
1969). 

In contrast, eleven Osprey eggs were found to 
contain more than 4 mg/kg DDE (Table 2). This 
concentration has been associated with 15% 
eggshell thinning in this species, and was proposed 
as the maximum level of DDE sustainable by a 
stable population (Wiemeyer et al. 1988). Two eggs 
collected in the Yukon contained mercury at levels 
within the hazardous range. 

Northern Harriers, Circus cyaneus, may have 
been affected by organochlorines prior to the 
1970s. Four eggs from the Prairies contained more 
than 10 mg/ kg DDE and one egg contained 5 mg/ 
kg dieldrin (Table 2). 

Falcons, which appear to be relatively 
susceptible to organochlorines (Fyfe et al. 1988) 



228 



The Canadian Field-Naturalist 



Vol. 104 



DDE (mg/kg) 




68 70 72 76 78 80 82 84 
YEAR 



86 88 90 



Dieldrin (mg/kg) 




68 70 72 76 78 80 82 84 
YEAR 



86 88 90 



Heptachlor epoxide (mg/kg) 



0.4 



03 



0.2 



0.1 { 



0.0 



I- 



9 



r 






g»0 



J 5 



i"'»"t t » 1^1 I » — I — I — I — r— I — I — r- 



68 70 72 76 78 80 82 84 
YEAR 

FiCiURf-: 2. Changes in residue levels in eggs of Swainson's Hawks collected in the Prairie provinces. 



86 88 90 



1990 



Noble and Elliot: Levels of Contaminants in Canadian Raptors 229 



were found to be highly contaminated (Appendix 
2D). Many MerUn eggs collected in the Prairies 
prior to 1973 contained DDE, dieldrin and 
heptachlor epoxide in excess of critical values 
(Table 2). By 1988, no eggs contained elevated 
levels of dieldrin, but DDE and heptachlor 
epoxide continued to be detected at relatively high 
concentrations (Table 2). Dieldrin levels were as 
high as 5.6 mg/kg, and heptachlor epoxide levels 
attained 9 mg/kg. Fox and Donald (1980) 
reported significant behavioural effects in Merlins 
where DDE concentrations in eggs exceeded 
8 mg/kg. Significant eggshell thinning is predicted 
to occur where the mean content of DDE in eggs 
exceeds 5 mg/kg (Fyfe et al. 1988). The Richard- 
son's race of Merlin which inhabits the Prairies 
feeds mainly on small flocking birds such as larks, 
longspurs and sparrows (Hodson 1978) although 
some individuals specialize in urban birds (Palmer 
1988). 

Analyses of a number of Prairie Falcon eggs 
collected between 1966 and 1988 in the same areas 
as the Merlins revealed relatively high concentra- 
tions of DDE, dieldrin, heptachlor epoxide and 
mercury (Appendix 2D). DDE concentrations as 
low as 1.2 mg/kg in eggs have been predicted to 
cause significant eggshell thinning, and to reduce 
productivity in this species (Fyfe et al. 1988). 
Prairie Falcons in Canada are mainly resident, and 
feed on a variety of small bird and mammals 
(Palmer 1988). 

DDE residues in all eggs of Gyrfalcons, Falco 
rusticolus were less than 15 mg/kg, but exceeded 
8 mg/kg in one egg. This arctic species feeds 
mainly on resident prey (Lincer et al. 1970). All 
American Kestrel eggs analyzed prior to 1973 
contained less than 10 mg/kg DDE, although one 
egg was high in dieldrin (1.25 mg/kg) and 
heptachlor epoxide (4.8 mg/kg) (Appendix 2D). 
During the late 1980s, mean DDE in five pools of 
Kestrel eggs from Ontario ranged from 0.44 to 
10.8 mg/kg, but levels of all other contaminants 
were low (Table 4; Appendix 2D). Kestrels feed 
mainly on invertebrate prey (Sherrod 1978), but 
often in agricultural areas with a relatively high use 
of pesticides. 

Many eggs of Peregrine Falcons contained DDE 
in excess of 15 mg/kg. The toxicological effects of 
DDE and other organochlorines on this species are 
discussed elsewhere in this issue (Peakall et al.) 

Owls were relatively uncontaminated (Appen- 
dix 2E), but we lack recent data. Only eggs of the 
Great Horned Owl were found to contain 
potentially hazardous levels of organochlorines, 
up to 16 mg/kg in a few samples. DDE 
concentrations of this magnitude have been 
associated with eggshell thinning in Barn Owls 
(Klaas et al. 1978). Two Great Horned Owl eggs 



also contained more than 1.5 mg/kg heptachlor 
epoxide (Table 2). 

None of the eggs of Snowy Owl, Nyctea 
scandiaca; Great Grey Owl, Strix nebulosa; Long- 
eared owl, Asio otus; Short-eared Owl, Asio 
flammeus or Burrowing Owl, Speotyto cunicula- 
ria, were found to have significant amounts of any 
contaminants. This is probably related to their 
basically mammalian diet (Godfrey 1986), a food 
source usually low in organochlorines (Lincer and 
Sherburne 1974). 

Temporal Changes 

Data permitting the evaluation of temporal 
trends in contaminant residues are Umited. Only 
eggs, which tend to be uniform in fat and water 
content, provide a standardized sample with which 
to compare among years. As the number of eggs 
collected at a particular location was insufficient, 
in most cases, to detect statistically significant 
differences, we have pooled eggs from different 
regions. Nevertheless, the extreme variability in 
contaminant levels in eggs, presumably due to 
individual and geographic differences in exposure, 
meant that few trends could be discerned. 

In the Prairies, contaminant residues in the eggs 
of four species were analyzed for temporal trends. 
Because earlier chemical analyses did not usually 
permit the identification of PCBs, chlordane 
metabolites, HCB or HCH, long-term trends are 
available only for DDE, dieldrin and heptachlor 
epoxide. 

In eggs of Swainson's Hawk from Saskatchewan 
and Alberta, dieldrin levels, but not DDE or 
heptachlor epoxide, have decUned significantly 
since the late 1960s (Figure 2). In eggs of 
Ferruginous Hawk, there were no significant 
declines in any of those three compounds {see 
Appendix 2B). 

In eggs of Prairie Falcons from Saskatchewan 
and Alberta, DDE, dieldrin, heptachlor epoxide, 
oxychlordane and PCBs generally declined 
between 1968 and 1988 {see Appendix 2D). HCB 
egg concentrations, however, increased during this 
time period. Merlins showed overall declines in 
DDE, dieldrin and heptachlor epoxide (Figure 3). 
Pairwise comparisons of residue levels found in 
viable eggs, revealed that DDE and heptachlor 
epoxide residues increased significantly between 
the earhest time period and the mid-1970s. 
Concentrations of all three compounds dechned 
significantly in the 1980s, although heptachlor 
epoxide levels did not differ significantly from 
those in the earliest collections. 

Very little long-term data are available for 
eastern Canada. If the analyses reported by Grier 
(1982) are included, it is possible to examine 
changes in contaminant levels in eggs of Bald 
Eagles from northern Ontario (Figure 4). Grier, 



230 



The Canadian Field-Naturalist 



Vol. 104 



DDE (mg/kg) 



Dieldrin (mg/kg) 




68 70 72 74 76 78 80 82 84 86 88 
YEAR 




68 70 72 74 76 78 80 82 84 86 88 
YEAR 



Heptachlor epoxide (mg/kg) 




■t T I I — r— I — I I f 

68 70 72 74 76 78 80 82 84 86 88 
YEAR 
FiCiURE 3. Changes in residue levels in eggs oi Merlins collected in the Prairie provinces. 



1990 Noble and Elliot: Levels of Contaminan rs in Canadian Raptors 



231 



DDE ('"S^S) 



Dieldrin (mg/kg) 



Mercury (mg/kg) 





. 










80 


- 




o 
o 






60 


- 




o 






40 


- 


o 


o 






20 



- 




•8c 


• 


t 



o CWS data 

♦ data from Grier (1982) 



1966 1968 1970 1972 1974 1976 1978 1980 1982 
YEAR 



3.0 
23 
2.0 
1.5 
1.0 
03 



0.0 



: 












- 




o 








1 










o CWS data 


" 










• data from Grier (1982) 


■ 


o 


8 








'■ 




o 
o o 


• 






' 




9 


• ♦ 


* 




\ 






• • 


• 




, 


1 


I.I. 


I.I.I 


. 1 . 





1966 1968 1970 1972 1974 1976 1978 1980 1982 
YEAR 





■ 






* 
• 


0.8 


o 








0.6 




o 




• 




0.4 


. 








OJ 


6 

o 


o 


• 
t 

• 


• 


on 


• till 


1 ■ 


1 . 1 ■ 1 


1 



o CWS data 

# data from Grier (1982) 



1966 1968 1970 1972 1974 1976 1978 1980 1982 

YEAR 

Figure 4. Changes in residue levels in eggs of Bald Eagles from northwestern Ontario. 



using some of the data reported here, found that trends. In eggs of Rough-legged Hawk, compari- 

DDE but not PCBs, dieldrin or mercury, declined sons between the earUest collections and those in 

in Bald Eagle eggs between 1970 and 1982. the 1980s, revealed no significant differences in 

Two arctic species were sampled at enough DDE, dieldrin or heptachlor epoxide. In eggs of 

locations to permit some analysis of temporal Gyrfalcons, no significant trends in any contami- 



232 



The Canadian Field-Naturalist 



Vol. 104 



Residues (mg/kg) 




Residues (mg/kg) 



1.50 
1.25 
1.00 
0.75 

0.50 
0.25 
0.00 



DIELDRIN 
HEPTACHLOR EPOXIDE 



5 1 



^ 



69 



I I I I t t I I' 

73 74 75 



80 81 83 



YEAR 

Figure 5. Changes in residue levels of organochlorine contaminants in eggs of Gryfalcons in northern Canada. 



nants could be discerned, although DDE, PCBs, 
heptachlor epoxide were highest in 1980 and 1981 
(Figure 5). 

There were insufficient data from Atlantic 
Canada or British Columbia to permit analyses of 
temporal trends. Although raptors have been used 
to monitor changes in environmental levels of 



persistent contaminants elsewhere (e.g. Froslie et 
al. 1986), the extreme variability in both eggs and 
tissue levels due to individual and geographic 
differences in diet, migratory behaviour and 
exposure, makes it difficult to discern trends. 
From the sparse temporal data available, we can 
conclude only that dieldrin and DDE occurred at 



1990 Noble and Elliot: Levels of Contaminants in Canadian Raptors 



233 



lower levels in Prairie raptors in the 1980s than 
during the early 1970s. Heptachlor epoxide levels 
have remained at low levels, except in a few 
individuals. 

The status of raptor populations in Canada 

The status of raptor populations in Canada is 
not well known, despite increasing interest in their 
welfare. The main sources of data on population 
trends come from counts along migration routes, 
Christmas Bird Counts, Breeding Bird Surveys, 
and research programs aimed at determining the 
status of particular species. 

Counts of most raptors are made annually 
during migration at a number of locations in 
North America, where movements of migrants are 
restricted by large bodies of water or mountain 
ranges (Heintzelman 1986). Of the more than one 
hundred such sites. Hawk Mountain, Pennsylva- 
nia, and Point Cape May, New Jersey, are the 
longest in operation. In the Great Lakes region, 
migrating raptors are monitored in Beamer and at 
Hawk Cliff (Ontario), Derby Hill (New York), 
and Duluth and Whitefish Point (Michigan). 
Data from these counts can be used to determine 
trends if certain assumptions (such as no overall 
changes in migratory paths) are made, and if 
weather and observer effort data are standardized 
(Sattler and Bart 1984). 

Long-term trends can also be estimated from 
Breeding Bird Surveys, an ongoing census of birds 
breeding along selected transects all over North 
America (Robbins et al. 1986). Other status 
assessments are derived from research employing 
a variety of census techniques ranging from aerial 
surveys of Bald Eagles (Gerrard and Ingraham 
1985) to detailed investigations of breeding 
success (e.g. Fox 1971). Nevertheless, the status of 
many species inhabiting remote areas, or whose 
nests are difficult to locate, has not been 
established. 

All methods of estimating numbers are subject 
to bias, and hence require cautious interpretation 
(Bock and Root 1981; Fuller and Mosher 1987). 
Table 3 summarizes the reported status of North 
American raptor populations, according to a 
number of reviews. Blue Lists are subjectively 
based on the opinions of experienced raptor 
enthusiasts from different regions (Tate 1986). 
Henny's (1972) assessments were based on the 
modelling of banding recovery data and estimated 
reproductive parameters. The results of counts of 
migrating raptors (Heintzelman 1986) and 
Breeding Bird Surveys (Robbins et al. 1986) are 
influenced by many factors, particularly weather 
and observer differences. However, as the biases 
inherent in the techniques employed to obtain 
these estimates are independent of each other, we 
feel that a consistent status assessment for a 



particular species can be considered to be 
accurate. 

We used the sources listed in Table 3, as well as 
the available literature on the status of particular 
species or populations (Noble and Elliott in press) 
in order to assign each species to one of four 
categories. 

The first group comprises species which are 
known to have experienced declines during the era 
of extensive pesticide use. It includes Bald Eagle 
populations in Ontario (Postupalsky 1971; Grier 
1982) and the Maritimes (Stocek and Pearce 
1978), Osprey in Ontario (Posupalsky 1972), 
Cooper's Hawk (Penak 1981), Merhns (Fox 1971; 
DeSmet 1985), Prairie Falcons (Fyfe et al. 1976) 
and Peregrine Falcons (Peakall 1976). 

Numbers of some of these species have 
recovered; others are still reduced. Gerrard and 
Ingraham (1985) noted that most Bald Eagle 
populations in Canada were stable or increasing, 
although they are still very rare in the Great 
Lakes. Ospreys apparently did not experience the 
dramatic declines noted in the United States, but 
in Nova Scotia, at least, it appears that 
productivity has increased since the mid 1970s 
(Fleming, personal communication). Although 
the extent and timing of population declines of 
Merlins in the Prairies has been debated (DeSmet 
1985), productivity and numbers do appear to 
have increased in recent years. Determining the 
effects of pesticides on Cooper's and Sharp- 
shinned Hawks is also problematic, as the 
evidence for declines in Canada is equivicol 
(Flood and Bortolotti 1986), despite the dramatic 
declines reported in the United States (Henny and 
Wight 1972; Snyder et al. 1973). Extensive 
shooting of these species in the 1950s and 1960s is 
one complicating factor; an ability to compensate 
for high mortality by increased productivity early 
in life is another possibility. 

The second category comprises species which 
are currently in decline, whether or not numbers 
are known to have declined in the past. Canadian 
raptors in this group include Red-shouldered 
Hawks (Risley 1983), Ferruginous Hawks 
(Schmutz and Schmutz 1980), Barn Owls 
(Campbell and Campbell 1983), Short-eared Owls 
and Burrowing Owls (Wedgewood 1978). 

The third category is comprised of species 
whose numbers have always been stable, or which 
appear to be increasing over most of their range. 
These include Red-tailed Hawks, Turkey Vultures 
and Great Horned Owls {see Table 3). 

The last category includes species for which 
there is no consistent evidence of either declines or 
increases (Table 3). These include most of the 
boreal forest owls. Golden Eagles, Northern 
Harriers, Gyrfalcons (Martin 1978), Northern 



234 



The Canadian Field-Naturalist 



Vol. 104 



Table 3. Status of North American raptor populations based on different sources. 



Source: 


Henny 1972 


Fyfe 1976 


Robbins et al. 1982 


Heintzelman 1986 


Blue List 


Time period: 


1946-68 


1975 


1965-79 


1961-84 


1980s 


Sharp-shinned Hawk 




stable 
decline in Ontario 


stable 


increase (1977-84) 


declining? 


Cooper's Hawk 


declining , 


stable 
decline in Ontario 


stable 


increase (1975-84) 


stable 


Northern Goshawk 




cyclic 




cyclic 




Red-tailed Hawk 


stable 


stable 


increasing 


stable 




Red-shouldered Hawk 


declining 


declining 


stable 


declining? 


declining 


Broad-winged Hawk 




stable 


stable 


stable 




Rough-legged Hawk 




fluctuating 




cyclic 




Ferruginous Hawk 




stable 






declining? 


Swainson's Hawk 




declining 






stable? 


Bald Eagle 




stable 

declining in 

Ontario 




increase (1975-84) 




Golden Eagle 




stable 




declining 


stable 


Osprey 


declining 


stable 


increase (east) 
declining (west) 


stable 


increase 


Northern Harrier 




fluctuating 
decline in Alberta 


declining 


stable 




Turkey Vulture 






stable 






Peregrine Falcon 




declining 




decHning (1972-84) 




Prairie Falcon 




stable in prairies 

decline in 
British Columbia 








Merlin 




stable 
decline in Ontario 




declining 




American Kestrel 


declining 
(1960-70) 


stable 

increase in 

British Columbia 




cyclic 




Gyrfalcon 




stable 








Great Horned Owl 




stable 
increase in Ontario 


stable 






Short-eared Owl 




fluctuating 






declining 


Long-eared Owl 




fluctuating 








Barn Owl 


stable? 








declining? 


Burrowing Owl 




stable 


declining 




declining? 


Great Grey Owl 




fluctuating 








Snowy Owl 




fluctuating 








Saw-whet Owl 




stable 









Goshawks, Rough-legged Hawks and Broad- 
winged Hawks, Buteo platypterus. Sharp-shinned 
Hawks were still blue-listed in some areas, but 
other populations were apparently healthy (Tate 
and Tate 1981; Tate 1986). 

There are many possible causes of raptor 
population declines, not the least of which is 
shooting, an activity that was prevalent until the 
1 960s. Other factors include loss of nesting habitat, 
reductions in prey abundance, disturbance of 
breeding sites, competition, and poisoning by 
currently used insecticides and rodenticides. 

Examination of the species in each category 
reveals some within-group similarities. Species in 



the first category which exhibited the most 
dramatic declines in the late 1960s, feed primarily 
on birds or fish. The relatively high levels of 
organochlorines found in migratory birds 
compared to mammals has been well documented 
(Eftderson and Berger 1970; Snyder et al. 1973). 
Since restrictions on the use of most organochlo- 
rine pesticides and mercurials in North America in 
the early 1970s, many of the species in this category 
have shown evidence of recovery (Grier 1982; Tate 
1 986; James et al. 1987). 

Although some species in the second category 
also feed on birds, most are species with relatively 
specialized habitat requirements. Reduced 



1990 



Noble and Elliot: Levels of Contaminants in Canadian Raptors 



235 



numbers of Red-shouldered Hawks in Ontario 
have been attributed mainly to loss of riparian 
deciduous forests (Risley 1983). In western 
Canada, the deterioration and loss of natural 
prairie and its replacement with crops and forage 
has undoubtedly contributed to declines in 
Ferruginous Hawks (Schmutz and Schmutz 1980) 
and the Burrowing Owl (Wedgewood 1978). 

Species in the third category tend to adapt well 
to human activities, profiting from new sources of 
food, or utilizing anthropogenic nest sites. Species 
such as the Red-tailed Hawk may have benefited 
by the reduced numbers of its competitors 
(Runyan 1988). 

The fourth category includes most of the arctic 
species, for which Uttle population data are 
available, and which have probably not been 
affected by changes in habitat. Numbers of 
Northern Harriers, Gyrfalcons and Rough-legged 
Hawks tend to fluctuate according to the 
abundance of their preferred prey; voles (Duncan 
1986), ptarmigan (Martin 1978) and lemmings 
(Godfrey 1986), respectively, and are therefore 
difficult to census. In species such as the American 
Kestrel, the negative impact of exposure to 
agricultural chemicals may have been mitigated by 
the increase in suitable open habitat and the 
increased availability of nest boxes. 

Conclusions 

Some Canadian raptors continue to be exposed 
to potentially toxic levels of organochlorines. 
Species such as Merlins, Peregrine Falcons, Sharp- 
shinned and Cooper's hawks which feed mainly on 
migratory birds are most contaminated. Rehable 
evidence of reproductive effects in Canadian 
raptors is restricted to a few species, namely Bald 
Eagles, Merlins, Peregrine Falcons and Prairie 
Falcons. However, high concentrations of DDE in 
eggs of those species as well as Osprey and Sharp- 
shinned and Cooper's hawks, were probably 
responsible for significant eggshell thinning and 
reduced productivity in the late 1960s and early 
1970s. Most owls and buteos. Golden Eagles, 
Gyrfalcons and American Kestrels are unlikely to 
have been affected by DDE. 

A few individuals and eggs of a number of 
species were found to contain significant 
concentrations of dieldrin, heptachlor epoxide or 
oxychlordane. There was no evidence that mercury 
has had any detrimental impact on Canadian 
raptors, although elevated concentrations were 
found in the livers of one Bald Eagle and several 
Turkey Vultures. 

Since the restrictions on the use of DDT and 
dieldrin were implemented in the early 1970s, 
concentrations detected in raptor eggs have 
generally declined. Populations of greatly reduced 



species (e.g. Bald Eagles in southern Ontario, 
Peregrine Falcons in eastern Canada) aided by 
reintroduction programs are showing signs of 
recovery, and there is some evidence to suggest that 
Merlins, Prairie Falcons, Osprey and Cooper's 
Hawk numbers have increased since the last 
decade. Declines in other species such as 
Ferruginous Hawks, Barn Owls and Red- 
shouldered Hawks have been reported more 
recently, and have been largely attributed to 
deterioration and loss of habitat. 

Nevertheless, the continued prevalence of 
organochlorine contaminants in the body tissues 
and eggs of Canadian raptors years after most of 
the parent compounds were banned suggests that 
we should continue to monitor them closely, 
particularly those feeding on migratory birds. 
Although we did not detect an effect of wintering 
area on contaminant concentrations, pesticide use 
in some Latin American countries where several 
Canadian raptors and their migratory prey species 
overwinter, may be a contributing source of 
current exposure. 

Acknowledgments 

Although too numerous to mention by name, we 
would like to acknowledge all of the people 
involved in the collection of this data; the field 
researchers who collected the samples, those who 
prepared the samples, the analytical teams, and 
those who created the databases. R. W. Fyfe, S. 
Postupalsky, J. W. Grier, G. A. Fox and N. 
Fimreite deserve special mention for their 
considerable contributions to the CWS data on 
contaminants in raptors. We also thank D. B. 
Peakall, W. K. Marshall and G. A. Fox for their 
comments on the manuscript, and A. Baril for 
extremely timely logistic support! 

Literature Cited 

Ames, P. L. 1966. DDT residues in the eggs of the osprey 
in the northeastern United States and their relation to 
nesting success. Journal of Applied Ecology 3 
(supplement): 87-97. 

Baril, A., J. E. Elliott, J. D. Somers, and G. Ericksen. 
1990. Residue levels of environmental contaminants in 
the prey of the Peregrine Falcon Falco peregrinus, in 
Canada. Canadian Field-Naturalist 104(2): 273-284. 

Bechard, M. 1981. DDT and hexachlorobenzene 
residues in southeastern Washington Swainson's Hawks 
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Received 6 February 1989 
Accepted 29 May 1990 



1990 



Noble and Elliot: Levels of Contaminants in Canadian Raptors 



239 



Appendix I A. Concentrations of organochlorines in tissues of Canadian raptors between 1967 and 1985. 





Prov. 


Tissue 


N 




Mean 


residue levels (mg/kg, wet weight) 




Species and year 


DDE 


Dieldrin 


HE 


Oxychlordane 


HCB 


PCBS 


Sharp-shinned Hawk 




















1969 


B.C. 


Brain 


1 


0.35 


0.043 










1982-85 


B.C. 


Livers 


7 


3.62 


0.128 


0.095 


0.17 


0.044 


3.51 


Cooper's Hawk 




















1967 


B.C. 


Brain 


1 


0.58 


0.013 










1984-85 


B.C. 


Livers 


3 


12.3 


0.63 


0.32 


0.48 


0.012 


4.77 


Red-tailed Hawk 




















1967 


Sask. 


Brain 


2 


0.068 


0.075 


0.016 








1968 . 


Alta. 


Brain 


1 


0.47 


0.15 


0.057 








1968 


B.C. 


Brain 


1 


0.39 


0.013 










Swainson's Hawk 




















1968 


Sask. 


Brain 


4 


0.085 


0.19 


0.131 








1968-71 


Alta. 


Brain 


5 


0.058 


0.041 


0.048 








Ferruginous Hawk 




















1969 


Sask. 


Brain 


1 


0.12 


0.013 


0.85 








1971 


Alta. 


Whole body 


2 


0.050 


0.027 


0.077 








Rough-legged Hawk 




















1973 


Yukon 


Brain 


3 


0.21 


0.014 


0.001 




0.010 


0.61 






Liver 


3 


0.074 


0.007 


0.002 




0.007 


0.21 


1974 


n.w.t. 


Liver 


I 


0.13 


0.020 


0.010 




0.020 


1.13 


Bald Eagle 




















1970 


N.B. 


Brain 


1 


7.09 


0.18 


0.062 




0.005 




1969 


Man. 


Brain 

Liver 


1 
1 


0.53 
2.12 


0.045 
0.11 


0.009 
0.017 








1968 


B.C. 


Brain 


1 


6.22 


0.10 










Golden Eagle 




















1973 


N.W.T. 


Brain 


2 


0.21 


0.010 


0.002 




0.007 


0.57 






Liver 


2 


0.28 


0.007 


0.001 




O.OIO 


0.54 


Osprey 




















1969 


Ont. 


Brain 
Liver 




0.22 
0.22 


0.013 
0.021 






0.017 
0.012 




1971 


Ont. 


Brain 

Liver 




0.10 
0.54 


0.020 










Northern Harrier 




















1968 


Sask. 


Brain 




0.039 


0.007 


0.030 








1968 


B.C. 


Brain 




0.55 


0.048 










Gyrfalcon 




















1968 


N.W.T. 


Liver 




0.040 


0.001 


0.001 




0.005 


0.14 


1971 


B.C. 


Brain 

Liver